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South African National
Heyday of the gymnosperms
systematics and biodiversity of the
Late Triassic Molteno fructifications
by
John M. Anderson and Heidi M. Anderson
i
Botanical
Pretoria
• •
T R E L I T Z I A
This series has replaced Memoirs of the Botanical Survey of South Africa and Annals of
Kirstenbosch Botanic Gardens which the NBI inherited from its predecessor organisations.
The genus Strelitzia occurs naturally in the eastern parts of southern Africa. It comprises
three arborescent species, known as wild bananas, and two acaulescent species, known as
crane flowers or bird-of-paradise flowers. The logo of the National Botanical Institute is
based on the striking inflorescence of Strelitzia reginae, a native of the Eastern Cape and
KwaZulu-Natal that has become a garden favourite worldwide. It symbolises the commit-
ment of the National Botanical Institute to promote the sustainable use, conservation,
appreciation and enjoyment of the exceptionally rich plant life of South Africa, for the ben-
efit of all its people.
Centenary Year
1 _ ^ ^
I In CELEBRATION
OF
THE CENTENARY OF THE NATIONAL HERBARIUM (PRE)* IN 2003 AND
ITS IMMEASURABLE CONTRIBUTIONS TO BOTANY IN SOUTH AFRICA AND FURTHER AFIELD
* See Bothalia 28: 271-297 (1998).
Cover design:
by Nadine Loots and Sandra Turck
incorporating drawings from this work
How to site this publication
ANDERSON, J.M. & ANDERSON, H.M. 2003. Heyday of the gymnosperms: systematics and
biodiversity of the Late Triassic Molteno fructifications. Strelitzia 15. National Botanical Institute,
Pretoria.
ISBN 1-919795-98-7
© Published by and obtainable from: National Botanical Institute, Private Bag X101, Pretoria, 0001
South Africa. Tel. +27 12 804-3200. Fax +27 12 804-3211. Website: www.nbi.ac.za. Typesetting and
layout by S.S. Brink (NBI). Printing by United Litho, PO. Box 40900, Arcadia, 0007 South Africa.
CONTENTS
FOREWORD iv
PREFACE v
ABSTRACT vi
GUIDE TO THE LAYOUT OF THIS VOLUME & SEQUELS . . vi
ON THE GENERAL SIGNIFICANCE OF THE MOLTENO vii
ACKNOWLEDGEMENTS
MOLTENO BIODIVERSITY & RELATED TOPICS
1. SAMPLING 2
2. FREQUENCY & ABUNDANCE 12
3. AFFILIATED ORGANS 16
4. MEASURING BIODIVERSITY 20
5. PROMINENCE (colonisation success) 26
6. THE MOLTENO BIOME 30
SYSTEMATICS of the MOLTENO GYMNOSPERMS
1. FORMAT OF THE SYSTEMATICS SECTION 42
2. CUTICLES 46
3. TAXONOMIC GUIDELINES 48
4. MORPHOLOGICAL TERMINOLOGY 49
5. CLASSIFIED LIST OF MOLTENO GYMNOSPERMS 54
6. ON THE PHYLOGENY OF THE PINOPSIDA 56
PINOPSIDA
Dordrechtites 60
Fredianthus 70
Lutanthus 74
Telemachus 82
Odyssianthus 88
Heidiphyllum 90
Clariphyllum 100
Rissikistrobus 102
Rissikianthus 108
Rissikia 112
Pagiophyllum 124
Gypsistrobus 126
Avistrobus 130
Helvetianthus 132
CYCADOPSIDA
Androstrobus 136
Pseudoctenis 140
Jeanjacquesia 142
Ctenis 144
Moltenia 146
GINKGOOPSIDA
Peltaspermum
Antevsia
Lepidopteris
Scytophyllum
Matatiella
Kurtziana
Switzianthus
De jersey a
Avatia
Eosteria
Ginkgoites
Paraginkgo
Hamshawvia
Stachyopitys
Sphenobaiera
Umkomasia
Pteruchus
Dicroidium
Fanerotheca
Kannaskoppia
Kannaskoppianthus
Kannaskoppifolia
Cetifructus
CLASS INCERTAE SEDIS
Alexia
Hlatimbia
Batiopteris
Hystricia
Saportaea
Linguifolium
BENNETTITOPSIDA
Fredlindia
Weltrichia
Cycadolepis
Leguminanthus
Halleyoctenis
Lindtheca
Taeniopteris
GNETOPSIDA
Nataligma
Gontriglossa
Graciliglossa
Cetiglossa
Fraxinopsis
Yabeiella
Jungites
UNDESCRIBED SEEDS .
BIBLIOGRAPHY
GLOSSARY
INDEX
148
154
156
170
172
176
182
186
192
194
198
208
210
216
222
240
250
256
272
286
290
294
314
318
322
324
330
332
334
336
340
341
342
344
356
358
362
364
368
369
370
376
384
386
390
396
398
iii
FOREWORD
Despite the bewildering diversity of living plants, it is re-
markable that the overwhelming majority are representative of a
single supremely successful clade: the flowering plants (angio-
sperms). These species range from grasses to baobabs, but as
descendants of a single common ancestor they share fundamental
similarities, and in a sense are merely different manifestations of
the same relatively recent evolutionary diversification. Where
flowering plants came from is a long-standing enigma in plant
evolutionary biology, but over the last two decades a key advance
has been the realisation that resolving the origin of flowering
plants — at least conceptually — is a straightforward issue. What
we need to know is: how are flowering plants phylogenetically
related to other groups of seed plants?
The greatest single obstacle to answering this question is our
ignorance of the diversity of other groups of seed plants (gymno-
sperms). Today, we have available to us only four non-
angiosperm groups — cycads, conifers, Gnetales and Ginkgo —
all of which are relatively well understood. In the past, however,
the palaeobotanical record shows clearly that there were once
many other gymnosperm groups that are now extinct. The in-
escapable conclusion is that the four groups of living seed plant
are a very poor and unrepresentative sample of all the different
kinds of seed plants that have ever existed.
This remarkable book on the gymnosperms of the Late Trias-
sic Molteno flora by John and Heidi Anderson is a major contri-
bution to understanding the diversity of seed plants that existed in
the past. It joins the earlier works by the same authors in reflect-
ing two lifetimes dedicated to exploring the biodiversity of the
past so as to better illuminate the biodiversity of the present.
Based on extensive fieldwork in the Karoo Basin of South
Africa, and huge collections (more than 27 000 catalogued slabs)
from almost 70 localities, this book provides the most compre-
hensive insight so far into extinct seed plants from the Mesozoic
of the Southern Hemisphere. Especially valuable is the use of
large numbers of specimens to delimit species, the careful note
taken of associations between different fossil species, and the
quantitative approach to sampling. I am not aware of any other
set of fossil assemblages that has been collected with such inten-
sity, such uniformity of approach and such care.
A key feature of this book is the focus on fossil seed plant
reproductive structures, which are represented by 35 genera and
more than 80 species. This component alone represents a huge
expansion to our knowledge of Mesozoic gymnosperms and pro-
vides the essential foundation for more detailed studies focused
on individual taxa in the future. Especially important are those
instances in which it has been possible to link ovulate structures,
pollen-producing organs and leaves. As detailed follow-up work
is done, these plants will become among the best-known plants
in the Mesozoic and they will have a significant impact on future
ideas of seed plant phylogeny and evolution.
Beyond its importance for systematic and phylogenetic studies,
this book paints a vivid picture of the exuberant diversity of life
in the Molteno flood plain, 230 million years ago. It reflects a
world quite different, and in many respects more modern, than
that of the Palaeozoic. It shows the re-emergence of biotic
diversity in the shadow of the Permian extinction, and docu-
ments the habitats in which modern groups such as mammals,
conifers, and beetles were beginning to come to the fore. But it
is still a world without flowering plants and many other modern
groups. As such it places our Recent condition in context and
provides important clues as to how the modern world came
about.
By any measure, John and Heidi Anderson’s dedication to
unlocking the secrets of the Molteno Formation has been extra-
ordinary and the result has been a landmark contribution to
palaeontology. This book and its previously published compan-
ion volumes will be of lasting value to anyone interested in the
biology and ecology of the Mesozoic. More generally, it pro-
vides an indispensable glimpse into a pre-flowering plant world
for anyone curious about plants of the past.
Peter R. Crane FRS
Director
Royal Botanic Gardens, Kew
Richmond, Surrey, TW9 3AB, UK
24 October 2002
IV
PREFACE
It was 35 years ago, in March 1967, that I ventured for the first
time into Molteno territory. From then till now the world has
changed. It has shifted — dramatically — in virtually every respect:
man has stood on the moon and looked back at this only known liv-
ing planet; we have come out the far end of the Cold War and of the
Apartheid era; and we have entered a new communications era, the
e-era, as overwhelming as that unleashed by Guttenberg’s printing
press in 1454. Most importantly of all, from our perspective, the
human population has doubled from 3 billion to 6 billion persons in
this time, and we have belatedly become aware of the Sixth
Extinction of life globally. In 1967 continental drift was only a
fringe hypothesis and concepts such as biodiversity and biophilia
were still well in the future. It is, indeed, only in the last decade
since the time of the First Earth Summit (Rio 1992) that the con-
servation fraternity has markedly expanded and begun expressing
severe concern at the decimation of life everywhere, across the con-
tinents and throughout the oceans. The world has indeed changed.
The falling of the Berlin Wall and the holding of the Rio Summit
virtually coincided. The political, economic and social energy, so
totally chained to the dread of nuclear annihilation can now be
focussed on a new, no lesser dread, the extinction of diversity
loosed on our planet by ourselves.
Back in the late 1960s when we began the long haul of amass-
ing close on 30 000 catalogued slabs from 100 Molteno tapho-
coenoses, we had no thought of biodiversity — nor did anyone else.
As the years rolled by and the number of localities discovered and
sampled grew, and as the number of fossiliferous slabs accumulat-
ed, I became simultaneously aware of the prodigious extant biodi-
versity in South Africa. Through a parallel project on indigenous
tree distributions I became woefully aware of the increasing pace of
destruction through human disregard of our biosphere. In searching
back and forth through geological and historical time, my aware-
ness of the vicissitudes in the wealth of nature grew synergistical-
ly. Living in the present time frame and working through past time
frames lends a sense of reality beyond oneself.
The Molteno became, increasingly, a lot more than merely an
exercise in collecting and describing fossils. It became for me far
more than a tally of papers published per year, far more than the
taxonomy of plants dead and buried 200 million-years ago. It
became part of a multifronted obsession to help swell our aware-
ness of biodiversity trends through time, and to impact somehow on
mankind’s collision with that diversity as it exists today.
This has meant spreading widely, perhaps too thinly. It has
meant, for one, that this volume on the Molteno gymnosperms—
Heyday of the gymnosperms — has taken a decade or two longer to
appear than it might have done. It has meant, also, a few stout
monographs instead of numerous slender papers in journals. So be
it; in our lives as lived in linear time, we can hope to have some
affect on the future, but not the past.
We have learned about ceaseless change and about ubiquitous inter-
dependence; about the many starts and the fewer enduring branches in
the tree of life— all of life— including our own individual lives. We are
one with nature. Here then, in this volume and its sequels, is the
Molteno gymnosperm story as currently known. Here is how it meshes
with and contributes to the broader picture of the evolution of life.
John M. Anderson
14 January 2002
ABSTRACT
A comprehensive study of the gymnospermous element of the Late Triassic Molteno Fm., Karoo Basin, South
Africa— based on a collection of 27 000 catalogued slabs from 100 assemblages (taphocoenoses)— is concluded
with the current volume whose focus has been very largely on the reproductive structures. This has revealed a
diverse flora including 20 genera (51 species) of ovulate fruit, 15 genera (35 species) of microsporangiate fruit
and 27 genera (113 species) of foliage, representing an estimated combined whole-plant total of 38 genera (143
species) in 32 families, 23 orders and 10 classes of gymnosperm. A high proportion of the female and male fruit
genera (and species) are described as new, while as many as 10 orders and 11 families are newly instituted. Of
particular interest are new families such as the Fredlindiaceae (based on the whole-plant Fredlindia/Halley-
octenis/Cycadolepis) and Lindthecaceae ( Lindtheca/Taeniopteris ), shedding new insight on the earliest evolu-
tionary radiation of the class Bennettitopsida; and the Antarctic order Petriellales (with Kannaskoppia/Kanna-
skoppifolia/Kannaskoppianthus from the Molteno) whose nearest affinities appear to lie with the Eurasian
Caytoniales. In spite of the relatively intensive and extensive sampling of the formation and the rigorous approach
towards identifying female/foliage/male affiliations, around half of all organ-genera remain without any sug-
gestion of affiliates and few are decisively established. Concluded from this is that the taphonomic filter is
severe, and that a significant proportion of the plant-genera colonising the Molteno Biome remain uncollect-
ed or were never preserved.
In order to place, systematically, the rich spectrum of new reproductive gymnosperm taxa found in the
Molteno, it has been necessary to prepare a revised global classification of the gymnosperms. This will be pub-
lished as sequels to the current volume. The classification is based, conceptually, on the ovulate fruit alone— with
a total of 8 classes, 37 orders and 76 families from the Late Devonian to present being recognised. The signifi-
cance with regard to Molteno systematics is the pattern of diversity trends that is revealed. At family and order
level there is seen a clear diversity low in the earliest Triassic and an equally clear peak (the 34 families being
twice the number recorded for any other interval) in the Late Triassic. It is from this evidence that we propose the
later Triassic as representing the heyday of the gymnosperms.
The Molteno Fm., as currently understood, provides the clearest available window globally onto the explo-
sive radiation of plant (and insect) life through the Triassic: it was the rapidly evolving ecosystems of this inter-
val following the end-Permian extinction that spawned the earliest mammals, dinosaurs and very possibly the
stem-angiosperms.
GUIDE TO THE LAYOUT OF THIS VOLUME & SEQUELS
The primary aims of this study (including its sequels) are to
describe the remarkable spread of gymnospermous fruit, many of
which are new and unique, discovered in the Molteno Formation; to
place these in context within a comprehensively revised classifica-
tion of the gymnosperms globally; and to draw attention to the peak
of biodiversity characterising the Late Triassic. The work is planned
to comprise two or three parts, with several sections, as follows.
THIS VOLUME
Molteno sampling, floristics & biodiversity
The purpose here is to provide a succinct backdrop to the sys-
tematics section. We present, largely in the form of tables, vital sta-
tistics on the 100 sampled taphocoenoses on the one hand and
results in terms of floristics, classification and diversity on the
other. The main thrust on Molteno biodiversity— with the far-
reaching insights this lends globally— is everywhere reflected.
Systematics of the Molteno gymnosperms
This section forms the core, and bulk, of the volume. Though
the emphasis is on the systematic description of the ovulate and
microsporangiate organs from the Molteno, the gymnospermous
foliage is also covered. A comprehensive account of the Molteno
gymnosperms, in classified sequence, is presented (see classified
list on pp. 54, 55). Introductory sections outline format and taxo-
nomic approach, and include a pictorial glossary of morphological
terminology. To lay stress on natural diversity, the affiliation of fruit
and foliage within the framework of the Molteno Biome, and its
seven recognised habitats, remains a guiding principle throughout.
Concise, comparative text accompanied by the liberal use of pen
sketches and photographic plates follows the style of our previous
monographs on the Molteno and other South African palaeofloras
(And. & And. 1983, 1985, 1989).
A total of 35 organ-genera (22 new) and 86 organ-species (79
new) of gymnospermous fruit, within an overall 32 whole-plant
families (11 newly named) and 23 whole-plant orders (10 newly
named), are described from the Molteno. (It must be emphasised
that families and orders are formally instituted and named here only
on the basis of ovulate — not microsporangiate or foliage — genera.)
SEQUELS TO CURRENT VOLUME
A Molteno overture
In this full-colour section, the purpose is to present an impres-
sionistic picture of the Molteno Biome: its climate, habitats (eco-
zones), flora, insect fauna, and, most particularly, the nature of the
gymnospermous element of the flora. The focus on the latter is
threefold: the diversity witnessed firstly amongst the female cones,
secondly amongst the male cones, and thirdly at whole-plant level
in distinguishing families and orders.
Whole plants: an exploration of habit & habitat
We have to date recognised 16 multi-organ genera in the gym-
nospermous fraction of the Molteno flora. Included are those cases
where affiliations between foliage and female and/or male fruits
have been established with some measure of reliability. Emphasis is
on Molteno distribution patterns, seasonality of production, and the
reference taphocoenoses— largely towards assessing the habit and
favoured habitat of the species differentiated within each genus.
Global classification of the gymnosperms
In view of the unsettled status of gymnosperm classification,
fossil and extant, we have felt compelled to attempt a global revi-
sion of the division to provide a meaningful context for the many
Molteno taxa. To enhance the integration between the Molteno and
global gymnosperms and for general usability, we have elected to
fully illustrate the classification. This compilation, including first
and last occurrences of all families, builds largely on Cleal (1993,
in The Fossil Record 2), the most recent, comprehensive classifica-
tion available, now a decade old.
vi
ON THE GENERAL SIGNIFICANCE OF THE MOLTENO
Of all the fossil plant/insect-bearing strata around the world, the
Late Triassic Molteno Formaton of the Karoo Basin, South Africa,
must rank as one of the most compelling. Resting immediately above
the celebrated bone-bearing, Permo-Triassic Beaufort Group-
second to none in revealing the story of the transition from reptiles
to mammals — lends it further interest. It has not yielded perminer-
alised fruit with exquisitely preserved anatomical structure as in the
coal balls of the Euramerican Carboniferous; or the consistent high-
quality cuticle of the Yorkshire Jurassic; or a multitude of fully
articulated insects as in a number of Cretaceous lagerstatte world-
wide; but for sheer potential in revealing the extraordinary richness
of the fecund later Triassic, it is apparently without peer. As cur-
rently known, it presents the clearest available window onto what
may well prove to be the most explosive of all plant radiations, that
leading to the acme of gymnosperm diversity and the ecosystems
spawning the mammals, dinosaurs and very possibly the stem-
angiosperms or flowering plants.
When we made our first collections from the Molteno 35 years
ago, there was no reason to suspect any of this. Little Switzerland,
with a superb view across to the Jurassic (flood basalts) escarpment
of the Natal Drakensberg, wonderfully picturesque itself between
sandstone cliff and wet indigenous forest, and with an obviously
diverse compression flora, offered a fine debut into the formation.
But only as we gradually, almost recklessly, added locality after
locality to the collection — and lobbied persistently for more storage
space — has the true significance of the Molteno come into focus.
Imagine a Michelangelo sculpture gaining progressively in defini-
tion as the marble slab is doggedly cleaved and chipped away.
But the most revealing aspect of the Molteno is what has not
yet been revealed, what we do not yet know about it, what remains
to be uncovered. The 100 sampled plant assemblages (tapho-
coenoses) with 27 000 catalogued slabs, 43 of which have yielded
insect faunas, almost literally represent just the tip of the iceberg.
Projections suggest that the 204 species of foliage and 333 species
of insect so far identified represent only a fraction of the total pre-
served flora and fauna. The Molteno exposure, the perimeter of a
tilted trapezium with dimensions of some 400 by 200 km, will evi-
dently yield hundreds more sites with further searching. According
to statistical extrapolations these promise to contain hundreds more
species, belonging to many still unknown genera, families and
orders of plant. And amongst this exuberant richness may well
appear some plants more angiosperm than gymnosperm.
The Molteno, in summary, proves uniquely engaging from a
range of interweaving perspectives.
The Triassic explosion. By statistical extrapolation, from observed
to preserved to existed species, the Molteno hints at biodiversity
figures for insects and plants apparently akin to those in the world
today. This strongly contradicts the conventional picture of a ‘ cone
of increasing diversity' through the past 450 million years. (And. &
And. 1995; Anderson et al. 1996)
From RNA interference to the Benny effect. With the ever-
increasing flood of discoveries, insights and hypotheses in molecu-
lar biology over the past decade in particular, we can look at inter-
vals of explosive evolution afresh. Scanning the pages of the most
ubiquitous fast-track scientific journals — Nature, Science,
Scientific American, New Scientist— of just the last year is literally
like peering into a new world. We need no longer confine our think-
ing to conventional Darwinian evolution through random mutation
and natural selection. The extraordinary renewal and radiation of
terrestrial life through the Triassic can be considered in the new
light of our deepening knowledge of the architecture and function-
ing of chromosomes, genomes and genes. Atavism, Hox genes,
stem cells, RNA interference, chromosome shuffling, gene transfer,
retroviruses, retroelements, inactive DNA, pre-programming, the
Benny effect, individually and together, offer new possibilities for
evolution within a period of dramatic radiation. Here we merely
cite these concepts; in the sequels to this work we probe their impli-
cations with regard to the Molteno flora, the Triassic explosion,
biodiversity patterns, gymnosperm classification and the gym-
nosperm heyday.
Heyday of the gymnosperms. The known Molteno fruit appear to
show at least 10 new orders of gymnosperm. A revised global clas-
sification (a sequel to this volume) of the division shows a clear
peak of gymnosperm diversification in the Late Triassic, with the
Molteno as the flagship formation. At the higher taxonomic levels
of order and class (basic morphological latitude), the gymnosperms
in this early-Mesozoic heyday seem to have outstripped the
angiosperms — with only two classes and some 84 orders (pre mole-
cular-based cladistics) — in their recent heyday.
Latitudinal diversity gradient. Wilson (1992), in The diversity of
life, defined a number of the generally acknowledged laws of bio-
diversity. One of the most clear-cut was the 'latitudinal diversity
gradient', of which he wrote: ‘[it is an] indisputable genera! fea-
ture of life that biodiversity rises towards the tropics.’
From our work on the Molteno and from an assessment of
diversity trends plotted against the Phanerozoic Ice-House/Hot-
House climatic curve (Scotese 1998; Scotese et al. 1999), we
offered the hypothesis (Anderson et al. 1999) that this rule might
hold, strictly, only for the relatively brief Ice-House intervals
occurring intermittently through geological time. One such unordi-
nary interval characterises today’s world. How well does the rule
hold up during the far longer Hot-House intervals when average
global temperatures rose to levels some 20°C hotter? 'In the Late
Triassic super-hothouse world at the time of the Molteno, for
instance, the definite possibility exists that a partial reversal of
today's latitudinal diversity gradient might have held. Energy lev-
els that are now optimal at the equator, might have been too
extreme during the Triassic ’ (Anderson 1999).
The world’s hottest terrestrial hotspot. Current evidence sug-
gests that the intriguing search for this Holy Grail of evolutionary
biology might well concentrate on the Late Triassic World — along
with the gymnosperm heyday. Further, in marked contrast to the
reality of our extant world, it seems to have centred on (southern)
temperate rather than tropical latitudes. The uplands bordering the
Molteno Floodplain Biome may feasibly lay claim to having sup-
ported the world’s richest terrestrial hotspot, past or present. We
offer this hypothesis as a challenge for debate.
The dawn of the extant world. It was approximately at the time of
the Molteno — providing the best available sample of the fecund
Late Triassic ecosystems— that the mammals, the dinosaurs, much
of the modern spectrum of insect orders (including the super-diver-
sity of the beetles and the abundance of cockroaches), and perhaps
the stem-angiosperms, all first evolved. The mammals and flower-
ing plants, both dominant today, seem to have followed a remark-
ably parallel ca 230 million-year life cycle, with the stem group of
each having arisen within the great Triassic explosion of life
(Anderson 1999).
It is particularly the third of these six themes, the heyday of the
gymnosperms, that we explore in this volume and its sequels. The
emphasis at all taxonomic levels is on biodiversity.
vii
ACKNOWLEDGEMENTS
Many persons have contributed in quite different ways towards making this volume on the Molteno possible.
We are grateful, firstly, for the continuing support of the National Botanical Institute, who have fully funded our
research since 1975.
Then there are the many farmers on whose lands we have excavated and through whose warm hospitality we
have thrived. Since our last volume in 1989, Fred and Linda Terblanche of Aasvoelberg, Isaac and Else Brummer
of Peninsula, and Piet and Sonya De Wet of Lutherskop have, in particular, welcomed us during our repeated inva-
sions from the city. Some 35 field trips have been specifically planned through the 1990s to improve our collec-
tions of reproductive material for this volume. Many of these were undertaken with Marijke, friend, and more
recently wife, of JMA.
Typing of the text has been done— somewhat fitfully over a good few years as the book steadily expanded—
particularly by Daleen Maree, Linda de Kock, Sheryl van Rooyen, and most recently Else van Doornum, Carrie-
Ann Greger and Nadine Loots. Nadine has also been largely responsible for scanning in the numerous sketches
and for the cover design. Sarie Brink has set the numerous tables and completed the final typesetting. Adela
Romanowski, as always, has willingly printed the photographs. Gill Condy and Nicolette Lavoyer (explicitly out
from Geneva) have helped with a number of the line drawings. Our daughter, Clara Anderson, and her companion
(now husband) Hannes du Plessis, have put the greater part of a month into recreating the 152 photographic plates
in Quark Express for electronic publishing. They were hijacked into this endeavour while on a break in South
Africa from their architectural careers in London.
A good number of palaeontological colleagues have offered critical comment on aspects of the manuscript at
various stages in its lengthy gestation. We particularly thank Chris Cleal, Wolfgang Meier, Andrew Scott, Greg
Retallack, Stephen McLoughlan, Raphael Herbst, Sid Ash, Conrad Labandeira and Hallie Sims in this regard.
Keith Holmes helped prodigiously in the sprint to the finish. We are deeply grateful to Peter Crane, Director of
the Royal Botanic Gardens, Kew, London, for so willingly writing the Foreword.
The Publications team, Louisa Liebenberg, Gerdt Germishuizen, Emsie du Plessis, Beverley Momberg, Sarie
Brink and Sandra Turck, at the National Botanical Institute here in Pretoria have put in a sterling effort towards
bringing our manuscript to reality in the Strelitzia series. We trust that their patience, stretched thin at times, will
be amply rewarded.
Finally, we wish to acknowledge the Ernest Oppenheimer Memorial Trust for generous sponsorship.
viii
^ lass#/ ..
OLTENO BIODIVERSITY
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& RELATED TOPICS
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d^TRELITZIA 15 (2003)
1. SAMPLING
Sampling strategy
Particular emphasis is given to the question of sampling in
view of its fundamental underpinning of any analysis of biodiver-
sity at successive taxonomic ranks. Two complementary sides to
sampling are separately considered.
Extensiveness of sampling : Extensiveness concerns the general
comprehensiveness of sampling, i.e. the total number of tapho-
coenoses (TCs, assemblages) covering the Molteno Fm. The even-
ness of spread across the geographic, lithostratigraphic, biostrati-
graphic and ecostratigraphic divisions of the formation is signifi-
cant. The frequency of occurrence of taxa through the Molteno is
thus documented.
Intensity of sampling-. Intensity concerns the particular compre-
hensiveness of sampling, i.e. the quantity of material examined
(measured in 'man-hours cleaving’) and curated (measured in the
number of ‘catalogued-slabs’ and/or ‘identifiable vegetative indi-
viduals’) for each TC. The relative or absolute abundance of taxa
within each TC is recorded.
The Molteno collection
The data recorded here account only for our own collections
housed at the National Botanical Institute, Pretoria (PRE/F/-) and
the Bernard Price Institute, Johannesburg (BP/2/-). Other limited
collections exist both in South Africa and overseas (see And. &
And. 1985).
Collecting programme'. 1967-1998 (31 years); 85 field trips.
Extensiveness of sampling: 100 taphocoenoses (assemblages) from
69 localities (areas to 1 km in diameter).
Intensity of sampling: ca 27 000 catalogued slabs including ca
300 000 identifiable vegetative specimens (individuals).
Geographic
grid (hierarchy)
Grid
diam.
Vol 1
1983
Vol 2
1989
This vol
2003
subregions
100 km
9
9
10
superlocalities
10 km
36
36
43
localities
1 km
44
48
69
sublocalities
100 m
-
-
80
supersites
10 m
50
57
85
TCs (assemblages)
Slabs (approx.)
-
67
13 400
75
16 600
100
27 000
Sampling history, 1967-1998
Sampling bias
In order to assess the comprehensiveness of sampling of the
Molteno Fm., four distinct perspectives are considered: geograph-
ic, lithostratigraphic, biostratigraphic and ecostratigraphic. How
evenly are the 100 Molteno TCs (69 localities, 43 superlocalities)
spread throughout the formation?
Geographic. For our Gondwana biogeographic maps we have
employed a degree-square grid (And. & And. 1983, 1989), a very
convenient scale when plotting distributions at the super-continental
level (Map 1, Tabs 2-4). To reflect biogeographic reality, all pro-
ductive grid squares — those yielding megaplant-bearing strata of
Triassic age — should be equally thoroughly sampled, an ideal far
from attained as yet.
The 10 degree squares covering the Molteno outcrop are
shown on Map 2, and the extensiveness of sampling within each
square is recorded in the accompanying Tab. 5.
The number of superlocalities per degree square varies largely
according to the thickness and area of outcrop. In degree square Ka
8, for instance, including the thickest Molteno sequences and with
as many as 1 1 productive superlocalities, the potential for discov-
ering further sites is great. The degree square to the West of Ka 1,
despite a considerable outcrop (though attenuated lithostratigra-
phy) and extensive searching, has, as yet, not yielded a single super-
locality.
Lithostratigraphic. The lithostratigraphic subdivision of the Mol-
teno into members— each consisting of a fining-upward cycle— has
not yet reached stability (Turner 1975; Christie 1981). We follow
here the six-member system and nomenclature of Christie (1981).
Our sampling varies significantly through the succession.
Cycle
Member
Sampling
6
Loskop
1 TC sampled
5
Tsomo
8 ”
4
Qiba
11 ”
3
Mayaputi
21 ”
2
Indwe
51 ”
1
Bamboesberg
8 ”
Lithostratigraphic sampling bias
The tally of TCs sampled per member is probably a fair reflec-
tion of potential. Half of all TCs are seen to occur in the Indwe
Member (Cycle 2), while only one is known from the Loskop
Member (Cycle 6), i.e. Kenegapoort (Ken 111 ) topping the list on
Tab. 1. The 100 TCs are placed with greater or lesser certainty
from oldest (in the Bamboesberg Member) to youngest (in the
Loskop Member) within the six-fold sequence.
A comprehensive lithostratigraphic and tectonic account of the
Molteno Formation and Karoo Basin can be found in Johnson etal.
(1997), Turner (1999) and Catuneanu et al. (1998).
Biostratigraphic. We provisionally recognise four assemblage
zones (Tab. 1 . left column) based on the combined megafloral and
faunal (insects) components of the formation (And. & And., in
prep.). These are not strongly differentiated and may better be
recognised as assemblage subzones, especially after systematic
cleaning of the ecostratigraphic imprint.
Assemblage zones (informal)
TCs
4
Sphenobaiera/Scytinoptera
9
3
Kannaskoppifolia sp.D/Ipsviciidae
32
2
Dicroidium zuberi/Moltenocupes
51
1
Kanaskoppifolia vincu/an'i/Permithonidae
8
Biostratigraphic sampling bias
Until further resolved, and for practical purposes, the biostrati-
graphic zones are taken to coincide loosely with lithostratigraphic
boundaries. The imbalance in sampling, as for the members, is
probably a reasonable reflection of overall potential.
Ecostratigraphic. Seven primary ecozones (habitats), illustrated
on p. 39, are recognised within the Molteno Floodplain Biome
(Caimcross et al. 1995). These are based on an overall analysis of
sedimentary strata, floral associations and insect faunas. They are
listed below from most to least complex.
Ecozones
TCs
1
Dicroidium riparian forest (mature)
2
2
” (immature)
8
3
” woodland
32
4
Sphenobaiera woodland
10
5
Heidiphyllum thicket
24
6
Equisetum marsh
18
7
Fem IKarmaskoppia meadow
4
-
uncertain
3
Ecostratigraphic sampling bias
The sampling is seen to be particularly uneven, with the
Dicroidium woodland (32 TCs) and Heidiphyllum thicket (24 TCs)
habitats together yielding over half the total TCs. The mature type
of Dicroidium riparian forest, with the richest plant assemblages, is
represented by only 2 TCs.
Sampling
g/tRELITZIA 15 (2003)
3
The Gondwana & Global Triassic
Gondwana Triassic (levels of sampling)
A measure of the undersampling of Gondwana Triassic strata
is documented in And. & And. (1983, pp. 2-27) and And. & And.
(1989, pp. 14, 15). By far the fullest succession of megafloras
(defined as deriving from a particular formation and basin) is pre-
served in the series of basins stretching down the eastern tectonic
margin of Australia. A full census of all Australian collections
made by one of us (JMA) in 1980 provides a quantitative basis for
comparison with Molteno sampling. Little collecting has been
done in the Australian Triassic since — aside from the Nymboida
Flora from the Middle Triassic Basin Creek Formation, Nymboida
Sub-basin (Holmes 2000, 2001).
Australia— 28 megafloras, 35 productive degree squares
203 ‘localities’, ca 300 TCs, ca 13 000 catalogued slabs
Molteno— 1 megaflora, 10 productive degree squares
36 localities, 100 TCs, ca 30 000 catalogued slabs
Australian Triassic & Molteno megaplant sampling
On a sampling scale of 1 to 5 (reconnaissance to comprehen-
sive), we might rate the Molteno grade 4 (approaching optimal)
and the Australian formations overall only 2 (basic) (And. & And.
Sampling intensity
grade
man-hours cleaving
1-2 3-9 10-49 50-100 >400
Reconnaissance
1
27
Low intensity
2
27
Med. intensity
3
33
High intensity
4
9
Top intensity
5
4
iii i
Sampling intensity grades
Optimal sampling: could aim at grade 3, 4 or 5 intensity
depending on the nature and diversity of the assem-
blage (TC)
1989, p. 15). Sampling of the Triassic outcrops around the other
Gondwana continents remains roughly on a par with Australia.
The near absence elsewhere in Gondwana of most of the repro-
ductive taxa described here from the Molteno will certainly to
some significant degree be a reflection of this undersampling.
Global Triassic (levels of sampling)
Triassic floras of the world overall remain highly unequally
sampled. Evidence of this is seen, for instance, in the massive im-
balances (Anderson et ai, in prep., sequel to this work) in the geo-
graphic origin of the type species of described and named ovulate
(gymnosperm) genera. Of the 58 genera now known from the global
Triassic, 17 are based on Molteno specimens, 21 on European
Triassic material, and only 13 from all remaining regions of the
Triassic world. (A further seven type species are of non-Triassic
origin.) It is hardly conceivable that these unequal numbers faith-
fully reflect phytogeographic or biostratigraphic significance. The
European bias is undoubtedly historical, while the Molteno bias
reflects, in significant measure, the level of sampling. It is abun-
dantly evident that, although the first ovulate strobilus (Triassic
gymnosperms) was described over 170 years ago (1828), we have
only begun to scratch the surface. We might safely predict a
notable increase in the unearthing and description of new genera,
representing new families and orders, in future years.
Geographic grid
Upper limit to areas
Name code
(hierarchy)
(maximum diameter)
(examples)
Region
1000 km
Saf(Ka)
Subregion
100 km
Ka (from Karoo Basin)
Superlocality
10 km
Lit (from Little Switzerland)
Locality
1 km
Lit 1
Sublocality
100 m
Lit II
Supersite
10 m
Lit III
Site
1 m
—
Subsite
10 cm
—
TC (assemblage)
10 m
Lit III Dic/Hei
The TC (taphocoenosis) is named after the dominant plants occurring in the
assemblage; in this case, Dicroidium 50% and Heidiphyllum 20%
Sampling hierarchy
Spacial occurrence: of fossiliferous strata (see And. & And. (1 983, p. 3) for full
details)
number of TCs
ro
o o
1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1
1 1«— 2-»
i
i
i
i
i
1
1
1
1
1
1
1
1 I I
1 1 1
' :
grade 4
1
1
! '
i
1
1
1
Iilll,
nIil, i, ,
,1
i
I
■ i
i
J
. i ; i '
i i
: : i
man-hrs — ► 10 20 30 40 50 60 70 80 90 100 400 550
cleaving
Fig. 4. Sampling intensity of the 100 Molteno assemblages (TCs)
Histogram: following this system of grading, around half the 100 Molteno TCs
have been sampled at the reconnaissance to low-intensity level and half at the
medium- to top-intensity level
Sampling
4
dA-RELITZIA 15 (2003)
SAMPLING
DIVERSITY
HABITATS
DOMINANTS
MOLTENO
Li
c
—
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catalogued
cleaving
if
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slabs
(man-hours)
species
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32
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14
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60
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21
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22
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116
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28
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184
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133
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1515
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1927
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120
550
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24
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141
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10
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63
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37
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98
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179
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124
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166
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13
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380
380
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23
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26
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60
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57
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589
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33
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88
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18
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339
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92
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104
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305
144
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1” 276
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22
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285
285
4
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/
78
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12
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55
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408
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152
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61
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53
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155
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288
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146
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462
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14
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157
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902
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400
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33
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303
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11
y
15
9 99 - 2
o
411 Dic/Sph
2176
2176
11
40
325
187
512
12
8
8
22
30
.
/
60 30 1 75 24
03
511 Die elo
27
27
1
1
2
2
11
11
V
50 20 20
<
Ask 111 Equ sp
161
205
366
2
3
12
2
14
1
1
7
5
12
/
21
1 - 50 9
m
Bam 111 Die dub
52
52
2
2
3
3
5
5
✓
98
- 1 - -
Total indivs (approx.)
16,600
10,600
27,200
2
8
32
10
24
18
4
3
Sampling
dv^TRELITZI A 15 (2003)
5
Sampling the primary habitats (Molteno)
Dicroidium riparian forest (mature, type 1) — Although this habitat
is the richest with regard to floral diversity, with Lit 111 (Little
Switzerland) yielding 38 vegetative species and Umk 1 1 1 (Um-
komaas Valley) 75 species, it remains the least extensively sampled
(with only two TCs). These sites are confined to the more norther-
ly outcrop area of the Molteno where the formation thins rapidly
and the potential for finding further TCs is relatively low. It is,
however, in this mature vegetation, interpreted as growing on the
earlier, erosive, Triassic landsurface — with the lowest Indwe
Sandstones of the Molteno unconformably overlying the Lower to
early-Middle Triassic Beaufort beds — where the greatest potential
evidently lies for significant new taxonomic finds. Umk 111, for
instance, has yielded three of the nine ovulate genera recorded, as
yet, from only a single TC.
Dicroidium riparian forest (immature, type 2) — This second cat-
egory of riverine forest (with eight TCs), interpreted to have never
reached great maturity owing to the shifting nature of the braided
rivers, is relatively rich in both vegetative species and ovulate
genera. The TCs are particularly varied and mixed in character in
that they were deposited under very different conditions on either
side of the narrow forested levees: either in near-bank channels of
the braided river or on the proximal floodplain associated with
crevasse splays. Sampling of this habitat is not particularly com-
prehensive and further focus on it will surely prove productive.
Dicroidium open woodland— By area, this vegetation type prob-
ably occupied a greater part of the Molteno Biome than any other.
It has been well sampled from 32 TCs. The floral diversity was
relatively low as is reflected in the low to medium diversity of the
preserved assemblages. Umkomasia and Peltaspermum are the most
frequently encountered ovulate genera, while other forms are few
and far between. There is evidently relatively little new to be dis-
covered.
Sphenobaiera closed woodland— It is these closed woodlands fring-
ing the lakes of the floodplain, along with the two grades of Dicroi-
dium riparian forest, that were the richest vegetation types of the
Molteno Biome. They have yielded the greatest diversity of ovulate
genera. With only 10 TCs, this habitat remains undersampled. The dis-
covery of further TCs of the quality of Bir 1 1 1 and Aas 41 1 will very
likely bring a good number of new fruit genera to light. It is pertinent
that Aas 4 1 1 has produced the record diversity of ovulate genera for
the Molteno (13 in all) and that three of these are unique to the site.
Heidiphyllum thicket— These low-diversity Heidiphyllum (conifer)-
dominated assemblages have been particularly well sampled from
24 TCs. They generally yield only two or three genera of ovulate
structure, most notably Telemachus (affiliated with Heidiphyllum),
Dordrechtites and Kannaskoppia. Other ovulate genera are rarely
encountered and further sampling of this habitat is unlikely to
unearth significant new reproductive finds.
Equisetum marsh— This vegetation type is often encountered in
the Molteno and, with 18 TCs sampled, has been well covered. The
original communities were presumably mostly monospecific and
the allochthonous gymnospermous elements in the fossil assem-
blages are rare: only two TCs of this category have yielded ovulate
or microsporangiate structures.
Fernl Kannaskoppia meadow— With only four TCs, these meadows
of the braid-river sandbanks remain undersampled. This is due to
the nature of the deposits: generally coarse, poorly bedded sedi-
ments reflecting rapid burial under turbulent flow conditions. Kan
1 1 1 and Kom 111, representing this habitat, are the only TCs from
the Molteno (aside from single Hamshawvia! Sphenobaiera and
Stachyopitys! Sphenobaiera specimens from Aas 41 1 and Maz 211
respectively; p. 16) that have yielded gymnospermous shoots with
fruit and foliage attached (see Kannaskoppia/ Kannaskoppifolia,
pp. 286-297). We anticipate that further sampling of this vegeta-
tion type may well yield important additional finds of fruit and
foliage in organic connection.
Annotated notes for TAB. 1.
Tab. 1. Sampling the Molteno Fm., 100 taphocoenoses (TCs)
Geographic: see Map 2 and Tab. 5
Lithostratigraphic: the full set of 100 sampled Molteno TCs is included in lithostratigraphic sequence according to the six
recognised sedimentary cycles (members); L=Loskop member (Cycle 6)
Biostratigraphic: the four provisionally defined assemblage subzones (manuscript in prep.) coincide approximately with the
lithostratigraphic boundaries as shown
Ecostratigraphic (habitats): it is indicated, for each of the 100 TCs, which of the seven primary habitats (ecozones) character-
ising the Molteno Biome is represented
Sampling: 27 200 catalogued slabs have been collected to date; 16 600 prior to our last monograph on the Molteno (And. &
And. 1989) and 10 600 subsequently
Diversity: the generic diversity of gymnospermous fruit and species diversity of all vegetative taxa are recorded
Dominants: to provide the essential character of each TC, the dominant gymnospermous genera and non-gymnospermous
groups are given
Bold = % estimate made at site
Mild = individuals in curated collection (where < 1%)
Sampling
6
d^TRELITZIA 15 (2003)
Tab. 2 GONDWANA TRIASSIC, PRODUCTIVE DEGREE SQUARES
Degree
o
ANTARCTICA (Ant)
square
>
o
<5
a>
TRANSANTARCTIC MTS. (TA)
SUPERREGION (continent)
£
_0) u
c
o
1 Allan Nunataks
76
155
3
4
-
2
1
3
REGION (basin)
ro
ra
~ 5
o
o
2 Taylor Glacier
77
155
3
4
3
1
2
9
Subregion (° square)
_i
o
O
O
O Q.
ffl
o
_l
3 Portal Mt.
78
155
2
3
-
1
1
1
4 Beardmore Gl.
84
160
1
2
3
SOUTH AMERICA (SAm)
5 Mt. Bumstead
85
170
2
4
_
2
1
1
CHILE (Ch)
6 Shackleton Gl.
85
180
2
4
_
1
1
1
1 Copiapo
27
70
3
3
- 2
1
3
EASTERN ANTARCTICA (EA)
2 Alto del Carmen
29
71
- 1
1
1
1 Prince Charles Mts
70
68
3
3
5
3
1
3
3 Los Vilos
32
72
2
2
- 2
1
2
4 Curico
34
72
- 1
1
1
AUSTRALASIA (Aus)
5 Pocillas
36
73
- 1
1
1
NEW ZEALAND (NZ)
6 Concepcion
37
73
3
3
- 2
2
2
1 SW Auckland
38
174
1
1
-
4
1
1
7 Temuco
38
73
- 1
1
2
2 Nelson Syncline
41
173
1
1
-
4
2
3
8 Valdivia
39
73
- 1
1
1
3 Mt. Potts
43
170
3
3
-
4
1
1
N. ARGENTINA (NA)
4 Benmore Dam
44
170
3
3
-
4
1
3
1 Ischigualasto
29
69
3
3
3 2
3
13
5 Southland Syncl. (NW)
45
168
2
1
-
4
5
10
2 Barreal
31
70
4
4
2 4
2
6
6 Southland Syncl. (SE)
46
169
2
1
-
4
2
7
3 Marayes
31
68
2
3
- 2
1
1
CANNING BASIN (Ca)
4 Cacheuta
32
70
4
4
3 3
3
10
1 Derby
17
123
2
3
2
2
1
2
5 Llantenes
35
69
3
3
- 3
2
2
2 Mt. Erskine
19
126
2
3
-
1
2
4
S. ARGENTINA (SA)
3 Mt. Ernest
20
126
2
2
-
1
2
8
1 Paso Flores
40
72
2
3
- 2
1
1
4 Gregory Lake
20
127
2
2
-
1
2
3
2 Los Menucos
40
69
1
2
- 1
1
1
SOUTH AUSTRALIA (SA)
3 El Tranquilo
47
70
2
3
- 2
1
1
1 Leigh Creek Cf.
30
138
3
3
5
3
1
1
PARANA BASIN (Pa)
2 Springfield Cf.
32
138
3
3
-
3
1
1
1 Santa Maria
29
54
2
3
- 2
1
1
GALILEE BASIN (Ga)
1 Lennox
22
146
1
2
1
1
3
SOUTHERN AFRICA (SAf)
BOWEN BASIN (Bo)
LUANGWA VALLEY (Lu)
1 Carborough
21
148
1
2
_
1
1
1
1 N. Luangwa V.
10
32
1
1
- 1
1
2 Springsure Shelf
24
147
1
2
_
1
1
1
2 W. Luano V.
14
29
- 1
1
3 Springsure Anticl.
24
148
1
2
_
1
1
1
ZAMBEZI VALLEY (Za)
4 Dawson Range
24
149
1
2
_
1
1
1
1 Kafue V.
15
28
- 1
1
5 Carnarvon Range
25
148
1
2
_
1
1
1
2 Urungwe
16
30
1
1
- 1
1
1
6 Taroom
25
149
1
2
_
1
1
3
3 Lake Kariba
17
27
3
3
- 2
1
8
CLARENCE/MORETON BASIN (CM)
4 Gwelo
19
29
2
2
- 2
1
1
1 Callide Cf.
24
150
2
4
5
2
1
1
LIMPOPO VALLEY (Li)
2 Gayndah
25
151
1
3
_
1
1
2
1 Soutpansberg
22
30
1
1
- 1
1
1
3 Tarong
26
151
2
2
2
1
1
5
2 Komatipoort Cf.
25
31
1
1
- 1
1
1
4 Esk Trough (N)
26
152
3
4
_
2
1
17
KAROO BASIN (Ka)
5 Ipswich/Esk
27
152
5
5
5
4
3
54
1 Bethlehem
28
28
1
3
5 2
1
1
6 Brisbane
27
153
4
5
4
3
2
14
2 Bergville
28
29
5
5
5 4
1
1
7 Nymboida
29
152
4
5
_
3
1
4
3 Maseru
29
27
5
5
3 4
1
4
8 Red Cliff
29
153
2
4
3
2
1
2
4 Underberg
29
29
5
5
3 4
1
9
NEW ENGLAND FOLDBELT (NE)
5 Aliwal North
30
26
5
4
- 4
2
13
1 Delungra
39
150
2
3
1
3
1
2
6 Zastron
30
27
4
4
- 4
1
2
2 Lome Basin
31
152
3
3
_
4
1
4
7 Matatiele
30
28
4
4
- 4
1
5
SYDNEY BASIN (Sy)
8 Molteno
31
26
5
4
- 4
1
19
1 Dubbo
32
148
3
4
_
2
1
2
9 Elliot
31
27
4
4
- 4
2
14
2 Blue Mts.
33
150
2
3
_
3
2
5
10 Maclear
31
28
4
4
- 4
1
1
3 Sydney
33
151
4
4
3
3
2
23
INDIA (Ind)
4 Picton
34
150
2
3
-
2
2
4
W. HIMALAYAS (WH)
VICTORIA (Vi)
1 Salt Range (W)
33
71
2
2
- 2
1
1
1 Yandolt Hill
36
143
1
1
-
2
1
1
PENINSULA INDIA (Pi)
2 Bald Hill
37
144
1
1
-
2
1
1
1 S. Rewa/Tiki
24
81
3
3
3 3
3
7
TASMANIA (Ta)
1 Poatina
41
146
1
1
1
2 Ben Lomond
41
147
3
4
3
2
1
2
22 regions
-
nil
3 NE coast
41
148
3
4
3
2
1
17
85 productive degree squares (subregions)
no information
4 Derwent Valley
42
146
3
4
3
2
1
2
94 biozones (megafloras)
5 Hobart
42
147
3
4
3
2
2
8
391 localities (69 Molteno, 322 non-Molteno)
6 South Cape
43
146
1
3
3
2
1
2
Sampling
(Sf ; TRELITZIA 15 (2003)
7
Tab. 3. GONDWANA TRIASSIC, STRATIGRAPHIC OCCURRENCE OF MEGAPLANTS
250 my
Updated from And. & And. (1983 & 1989)
Tab. 4. THE MEGAPLANT-BEARING FORMATIONS OF THE GONDWANA TRIASSIC
SOUTH AMERICA
Chile (Ch)
34 La Ternera
” El Puquen
" Gomero
” Tralcan
33 Gomero
” Tralcan
25 Quilacoya
24 Quilacoya
18 (Alto Del Carmen)
N. Argentina (NA)
28 Ischigualasto
” Q de la Mina
27 C. de Piedra
25 L. Carrizal
" Cacheuta
24 Panul
” Potrerillos
” Llantenes
23 Los Rastros
” Cortaderita
" Chihuiu (U)
22 Ischichuca
” Barreal
18 Las Cabras (M/U)
17 Las Cabras (L)
S. Argentina (SA)
24 Paso Flores
23 El Tranquilo
22 Los Menucos
Parana Basin (Pa)
28 Santa Maria
SOUTHERN AFRICA
Luangwa Valley (Lu)
24 ‘U. Grit'
12 Ntawere
Zambezi Valley (Za)
24 ‘Flags’
Limpopo Valley (Li)
24 (Molteno)
Karoo Basin (Ka)
25 Molteno
24 Molteno
12 Burgersdrop
INDIA
W. Himalayas (WH)
12 Landa
Peninsular India (PI)
29 Tiki
24 Chicharia
12 Parsora
ANTARCTICA (TA, EA)
Transantarctic Mts.
22 Falla
” Lashly C
19 Fremouw
10 Lashly A
Eastern Antarctica
29 Flagstone Bench
AUSTRALASIA
New Zealand (NZ)
34 (Southland)
28 (Southland)
21 Tank Gully
” Black Jacks
” Long Guily
20,16,14 (Southland)
13,11,9 ( " )
Canning Basin (Ca)
11 Culvida
9 Erskine
7 Blina
South Australia (SA)
22 Leigh Creek
” Springfield
Galilee Basin (Ga)
18 Moolayember
17 Clematis
Bowen Basin (Bo)
19 Moolayember
17 Clematis
Clarence/Moreton Basin
35 Raceview
34 Aberdare
24 Callide
” Ipswich (U)
23 Tarong
’’ Ipswich (L)
” Tingalpa
” Red Cliff
19 Esk
” Nymboida
18 Neara
17 Bryden
New England Foldbelt
20 Gragin
19 Gunnee
9 Camden Head
Sydney Basin (Sy)
18 Benolong
" Wianamatta
17 Hawkesbury
11 Burralow
” Gosford (U)
” Newport (U)
10 Newport (M)
9 Garie
” Newport (L)
8 Banks Wall
" Gosford (L)
” Bald Hill
7 Patonga
” Bulgo
Victoria (Vi)
22 Yandoit
’’ Bald Hill
Tasmania (Ta)
25 Brady
” New Town
7 Knocklofty
Updated from And. & And. (1989)
Explanatory notes
(a) The table provides the base for showing the stratigraphic distribution of each genus.
(b) In the interest of ready comparison with our previous Molteno monographs (And. & And. 1983, 1989), no attempt has been
made to update correlations or to introduce the latest standard time scale. A series of fully annotated correlation charts appears
in And. & And. (1983).
(c) N — Norian L — Ladinian S — Spathian D — Dienerian
C — Carnian A — Anisian S — Smithian G — Griesbachian
(d) The 35 units into which the Triassic is subdivided represent the international standard ammonite biozones. In that the dura-
tion of the Triassic Period (see p. 45) was about 47 million years (250-203 my), each unit or zone represents about 1 ,3 million
years.
(e) The 22 Gondwana regions (’basins’) yielding Triassic megaplants are each allotted a single column. Antarctica is an excep-
tion: Eastern Antarctica (Flagstone Bench Fm. of the Prince Charles Mts.) has been newly incorporated in the table but not
given a separate column — A* = TA + EA.
(f) i 1 The coloured bars represent megaplant-bearing strata. For convenience each productive formation (or other sets of
beds) is taken to coincide in duration with a particular ammonite biozone.
(g) A bar may represent a single productive formation or a number of correlative formations in adjacent subregions (degree
squares). In a few instances (e.g. Molteno, Ipswich) a formation is represented by 2 bars.
(h) The productive ’formations’ are listed youngest to oldest for each region and numbered according to the ammonite zonation.
Sampling
^/tRELITZIA 15 (2003)
w
Sampling
^/tRELITZIA 15 (2003)
9
Degree
square
Superlocality
Super- Assem-
site blage
Supersite
(informal)
Degree
square
Superlocality
Super- Assem-
site blage
Supersite
(informal)
Ka 2
1
Little Switzerland
Lit 111 Dic/Hei
-
1
22
Birds River
Bir 111 Sph 2spp
Tenant Dam
[
k
2
Champagne Castle
Cha 111 Die odo
Champagne Hotel
.. 211 ! ’
Cloete Shale
” 211 Die dub
Wonder Valley
” 311 Hei/Sph
van Biljoen Shale
3
Injasuti Valley
Inj 111 Die odo
Injasuti Camp
23
Bamboesberg
Bam 111 Die dub
-
" 211 Die dub
Forest Patch
24
Aasvoelberg
Aas 111 Hei elo
Turner Shale
4
Mooi River
Moo 111 Die zub
-
’’ 211 ” "
Proppie Shale
Ka 4
5
Hlatimbe Valley
Hla 111 .Equsp
Homestead Boulder
"311 " ”
Easter-Egg Shale
” 211 Die 3spp
Tom’s Outcrop
"
” 411 Dic/Sph
Fredlindia Slate
” 212 ; "
Eddie's Outcrop
" 511 Die elo
Erard Slate
” 213 i Die elo
Batiopteris Corner
" 611 Hei elo
Aasvoelberg Pass
'
1
6
Umkomaas Valley
Umk 111 Die 2spp
Waterfall Locality
25
Boesmanshoek Pass
Boe 111 Lep sto
Equisetum Ridge
7
Sani Pass
San 111 Die era
-
” ” Equ sp
8
Mngeni Valley
Mng 111 Die 2spp
-
Ka 8
” " Dic/Hei
9
Qachasnek
Qac 111 Hei/Dic
-
" 112 Die cor
Coriaceum Flats
10
Matatiele
Mat 111 Die dub
-
26
Cyphergat
Cyp 111 Die era
Open Cast
Ka 7
11
Kenegapoort
Ken 111 Die era
-
" ” Hei elo
-
12
Tina Bridge
Tin 111 \ Sch sp
Tina Road
27
Molteno
Mol 111 Sph pon
Slate Quarry
” 121 ; Sph 2spp
Tina Quarry
28
Kleinhoek
Kle 111 Equ sp
Kleinhoek Cutting
” 131 Hei/Ast
Tina Stream
" ” Hei elo
Ka 10
13
Waldeck
Wal 111 Die odo
" ” Hei/Dic
i
k
14
Konings Kroon
Kon 111 i Die odo
Du Toit Chert
" 211 ; Sph pon
Kleinhoek Slate
" " ; Sch sp
29
Kapokkraal
Kap 111 Dic/Ris
-
" " Hei elo
30
Kommandantskop
Korn 111 Sph/Dic
-
” 211 Ast2spp
Rooipoort Donga
31
Kullfontein
Kul 111 : Sph pon
-
” " Hei elo
J
32
Telemachus Spruit
Tel 111 Hei elo
-
” 221 Ast 2spp
Stream
i
[
33
Vineyard
Vin 111 Die odo
Homestead
" 222 Die odo
Rosy Chert
” 211 1 Sch sp
Cheese Factory
" 223 ; " ”
Keith Chert
34
Elandspruit
Ela 111 Die odo
Anne Roadside
15
Peninsula
Pen 321 Dic/Ris
Rissika Chert
” 112 Equ sp
Zorba Stream
" 211 Dic/Equ
Isaac Chert
” " Dic/Hei
„ 22 1 - »
Volker Chert
35
Kraai River
Kra 111 Die odo
Jigsaw Chert
” 511 ] Equsp
Mudpatch Chert
” 211 Equ sp
Plaatkop Track
Ka
9
" 421! Die odo
360° Chert
” 221 beetles
Plaatkop Beetles
" 431; Dic/Equ
180° Chert
” 311 Die odo
Glenburn
" 311 'Hei elo
Campsite Quarry
Ka 5
36
Lutherskop
Lut 111 Hei/Dic
Potgieter’s Place
" 411: " "
Lunchspot
1
'
” 211 Equ sp
Turner's Tributary (1)
16
Kannaskop
Kan 111 Ast spA
Kannaskoppia Siltstone
” 221 | ” ”
' (2)
" 112’ Hei elo
Upper End
" 311 Hei elo
Oom-Piet's Campsite
17
Navar
Nav 111 1 Equ sp
Elliot Quarry
" 4111 Equ sp
Orange View
" " Die odo
" 4112 Hei/Dic
18
Cala Road
Cal 111 Equ sp
Cala Cutting
" 511 Hei elo
Sonya's Hillock
" " Dic/Sph
37
Nuwejaarspruit
Nuw 111 Equ sp
Clara's Ditch
" 211 Hei elo
Cala Bolders
" ” Die zub
19
Askeaton
Ask 111 Equ sp
-
" 211 Die 2spp
Wolfy's Adit
20
Greenvale
Gre 111 1 Sph pon
Equisetum cutting
Ka 6
38
Winnaarspruit
Win 111 Hei elo
-
" ” Equ sp
39
Qualasi Hill
Qua 111 Die odo
-
” 121 Hei elo
Telemachus Shale
t
Ka 3
40
Morija
Morlll Die zub
-
21
Dordrecht
Dor 111 Hei elo
-
" ” 1 Die odo
-
41
Makoaneng
Mak 111 ! " "
-
l
42
Mazenod
Maz 111 Die era
Luka
" 211 Hei/Dic
Lechesa
Ka 1
43
Golden Gate
Gol 111 Die dub
-
Tab. 5. Molteno ‘localities’ and assemblages
Superlocalities', numbered in clockwise order around the Molteno outcrop Extensiveness of sampling
(see map opposite) 10 degree squares/subregions (ca 100 km diam. grid)
Supersites : arranged chronologically, within each superlocality, according to date 43 superlocalities (10 km diam. grid)
of discovery (earliest above) 69 localities (1 km diam. grid)
Assemblages', arranged stratigraphically for each supersite (youngest above) 80 sublocalities (100 m diam. grid)
Informal names: all distinct supersites within a superlocality have been given a 85 supersites (10 m diam. grid)
familiar name for ease of communication (assemblages within a 100 assemblages (taphocoenoses, TCs)
supersite retain only the formal code name)
Tab. 6a, b (see overpage). Molteno gymnosperms, vegetative & reproductive genera (abundance & frequency)
Assemblages (taphocoenoses. TCs): the full set of 100 sampled TCs is arranged in lithostratigraphic order following the 6
recognized sedimentary cycles (& members)
Gymnosperm genera: the 27 foliage genera, 20 ovulate genera & 15 microsporangiate genera are listed in classified
sequence
Productive TCs (reproductive genera): 57 of the 100 TCs yield ovulate &/or microsporangiate taxa
Abundance: the relative (% in bold) or absolute (individuals in mild) abundance of genera per TC is recorded within the
matrix of the table
Sampling
10
d^TRELITZIA 15 (2003)
Molteno members
Molteno cycles
Tab. 6a.
MOLTENO
FOLIAGE
assemblage
(taphocoenosis)
NON-
GYMNOSPERMS
GYMNOSPERMS
generic
diversity
■2
3
§
Marchantites
Thallites etc.
lycopods
horsetails
| ferns
Pinopsida
| |
•c .2
.9. o. as
!2 'c 35
55 -2 -!2
a: 0 o:
| Pagiophyllum
Pseudoctenis
Cycadop.
•2
Q>
a
o*
.TO y>
TO |
3 O
| Moltenia
Q)
Q.
■§
1
Scytophyllum
Kurtziana
9.
Dejerseya
(O
Ginkgoites 0
T3
Paraginkgo 5;
0)
Sphenobaiera
Dicroidium
| Kannaskoppifolia
B adopter is
3
Saportaea g-
| Linguifolium
Halleyoctenis
3
Taeniopteris ?
Gontriglossa
Gnetopsida
TO
V)
•2 $ TO
if!
0 0 £
Jungites
i
a
0
c
£
>.
03
c
0
c
gymnosperms
total
J 1 6
Ken 111
Die era
1
3
99
2
1
3
Nav 111
Equ sp
100
1
1
Die odo
1
98
1
3
3
2 '
Cal 211
Hei elo
20
75
5
1
2
3
o' s
" 111
99
1
2
2
<0 1 5
Dic/Sph
1
3
44
5<T
1
3
4
Bir 211
Sph 2spp
1
1
3
3
95
4
2
2
5
7
" 311
Hei/Sph
45
1
40
14
1
5
5
1
" 111
Sph 2spp
40
4
7
4
20
90
10
1
1
70
43
85
1
1
6
70
40
11
11
22
1
Dor 111
Hei elo
5
85
10
1
2
3
1
Gre 121
2
20
2
98
~2~
3
22
3
4
7
" 111
Sph pon
10
5
85
3
3
Equ sp
1
97
i
1
2
1
1
4
4
8
Boe 111
Lepsto
2
7
90
5
3
5
2
5
7
CQ I 4/5
Egusp
100
1
1
O l
Dic/Hei
42
7
57
r ■
1
5
5
" 112
Dicca
4
14
12
3
99
1
1
5
6
Cyp 111
Die era
6
1
1
4
24
75
83
20
2
4
5
9
1
Hei elo
100
1
1
l
Md 111
Sph pon
99
12
2
2
i
Kan 112
Hei elo
6
98
T
19
2
1
4
5
9
" 111
AstspA
22
63
10
2
5
2
3
5
Tel 111
Hei elo
i
48
58
89
1
23
6
33
1
2
6
7
13
1
Kom 111
Sph/Dic
1
15
1
60
39
30
3
3
6
1
Vin 111
Die odo
1
2
28
3
7
1
4
70
2
4
2
8
10
1
Elalll
4
7
2
5
73
V
87
T"
6
1
8
9
Kra 311
5
99
18
3
3
" 211
Equsp
100
1
1
" 221
beetles
50
1
50
2
1
3
2 1 ,
" 111
Die odo
1
4
10
4
1
90
5
3
5
8
Lut 111
Hei/Dic
1U
50
40
1
2
3
< |
" 511
Hei elo
1
80
20
1
2
3
" 4112
Hei/Dic
50
50
2
2
1
" 4111
Equsp
100
1
1
1
" 311
Hei elo
6
1
30
3
99
19
5
29
58
66
23
9
4
8
12
1
" 221
Equ sp
99'
1
1
1
2
" 211
80
2
10
10
4
2
3
5
1
Tin 121
Sph 2spp
1
4
1
1
95
1
2
1
1
7
8
1
" 111
Sch sp
89
10
2
1
3
1
" 131
Hei/Ast
10
40
50
2
1
3
1 2/3
Wallll
Die odo
2
1
2
"2
"3"
92~
3
20
8
8
Kon 223
1
1
4
4
4
1
80
1
15
1
5
9
14
" 222
2
1
1
9
3
9
13
10
87
8
4
1
7
9
16
1
" 221
Ast 2spp
1
20
52
25
2
2
2
1
7
5
12
1
" 211
20
52
21
4
6
10
10
3
6
6
12
" 211
Hei elo
9b
5
2
2
" 111
Die odo
1
1
12
7
2
5
24
5
78
10
1
1
3
9
12
1
Sch sp
90
10
1
1
2
l
Hei elo
10
84
5
3
1
3
4
l
Pen 321
Dic/Ris
3?
5
1
35
1
9
5
51
5
5
2
4
9
13
| f
" 211
Dic/Equ
1
45
2
'2
1
50
2
4
4
8
" 221
40
20
2
2
40
3
3
6
l
" 511
Equ sp
98
2
1
1
2
l
” 421
Die odo
5
4
5
1
4
89
2
1
6
7
1
" 431
Dic/Equ
40
5
5
40
2
2
4
6
" 311
Hei eilo
6
2
5
11
75
25"
41
5
3
8
" 411
1
3
2
3
94
50
13
80
6
4
10
1
Weill
Equsp
99
1
1
1
2
1
Hei elo
10
90
1
1
2
l
Hei/Dic
49
3
1
8
45
5
6
6
" 211
Sph pon
TOO
1
1
1 e
Kul 111
2
1
99
1
2
2
4
-,1
Kap 111
Dic/Ris
10
4
25
30
38
40
2
20
50
6
4
3
1
5
3
12
15
£!• 1
Vin 211
Sch sp
100
1
1
g!
Ela 112
Equ sp
95
4
1
2
1
3
Dic/Hei
10
lO"
60~
1
2
3
- , d
Nuw 111
Equsp
i
99
2
2
i
Diczub
4
2
30
70
1
1
1
5
6
i
" 211
Dic2spp
1
1
3
98
4
4
Win 111
Hei elo
1
2
3
79
10
10
4
1
5
5
10
Main
Diczub
1
99
T
1
2
3
Die odo
1
98
2
1
2
3
Qua 111
20
20
40
5
4
4
Mak 111
7
1
90
10
4
4
i
Maz 111
Die era
5
3
5
2
1
12
74
1
22
3
7
10
" 211
2
7
4
32
1
1
3
1
■64"
1
T
11
4
9
13
Moo 111
Diczub
5
99
2
2
i
Hla 111
Equ sp
99
1
1
1
2
i
" 211
Die 3spp
10
1
1
3
2
1
7
85
2
1
3
8
11
" 212
1
6
1
1
4
1
7
91
1
5
1
2
9
11
" 213
Die elo
3
2
1
3
11
32
T
2"
27
7
' V
1
49
89
7
"2"
6
8
1
16
13
29
Umk 111
Dic2 spp
41
26
2
i
7
20
5
1
1
41
1
1
18
5
69
42
2
55
5
13
3
45
19
18
37
Chain
Die odo
100
1
1
" 211
Die dub
20
80
1
1
2
Inj 111
Die odo
1
100
4
2
1
3
4
" 211
Die dub
70
"90"
1
3
3
i
San 111
Die era
1
2
1
5
1
1
1
5
3
90
3
2
2
3
10
13
i
Mng 111
Die 2spp
6
2
2
6
93
2
6
6
i
Qac 111
Hei/Dic
1
2
50
8
50
2
3
5
Matlll
1
1
1
20
7
4
1?
1
3
25
18
89
2
2
2
9
10
19
Gd 111
2
Lit 111
Dic/Hei
14
2
10
10
23
40
48
6
1
1
20
4
40
1
50
56
1
17
8
78
30
6
18
6
19
25
i
Aas611
Hei elo
1
80
20
1
2
3
Si
" 111
17
10
20
77
1
7
2
2
6
5
11
" 211
10
6
2
99
14
19
3
3
6
" 311
18
13
2
99
1
9
15
26
1
4
6
10
" 411
Dic/Sph
5
3
50
75
24
1
25
2
19
7
15
41
30
60
150
40
1
75
7
13
20
CO 1
" 511
Die elo
20
10
20
50
1
5
5
Ask 111
Equsp
1
50
9
20
1
21
6
3
9
CO 1
1
Bam 111
Die dub
1
1
1
98
4
4
Frequency; Total TCs
18
18
6
7
60
56
62
3
21
1
21
3
2
5
30
1
13
5
19
2
43
75
26
10
1
9
10
38
8
1
1
29
1
Tab. 6a. Molteno gymnosperms, foliage genera (abundance & frequency)
see p. 9 for notes
Sampling
^/tRELITZIA 15 (2003)
Tab. 6b.
MOLTENO
9 & cf
fruit
assemblage
(taphocoenosis)
Ovulate genera
Pinopsida
Ginkgoopsida
- =2 >
Microsporangiate genera
Pinopsida
£ -c
S c
•2 .2
Cy.
Ginkgoopsida
a a •§ g c
i 8 a s S
co uj to a. ic
Die era
Equ sp
Dicodo
Hei elo
Db/Sph
Sph 2spp
Hei/Sph
Sph 2spp
Hei eta
Sph pon
Equsp
Lepsto
_Egu sp_ _
111
Tel 111
Kom 111
Vin 111
Elalll
Kra 311
211
221
. 111_ _
Lut 111
" 511
" 4112
" 4111
" 311
221
211
Kul 111
. Kap 111
Vin 211
Ela11_2_
Nuwlll
Qua 111
Mak 111
Mazlll
" " 211 "
Moo 111
Hla 111
" 211
"_212_
”” 213
Umk 111
Chain
211
lnj_111 _
"211
San 111
Mng 111
Qac 111
Man 11 _
Gol 111
□till
Aas611
111
211
Dic/Hei
Die cor
Die era
Hei eb
Sph pon
Hei eb
AstspA
Heieb
Sph/Db
Dbodo
Equsp
beetles
_Dicqdo_ _
Hei/Db
Heieb
Hei/Db
Equ sp
JHet eb
Equ sp
Sph 2spp
Sch sp
Hei/Ast
Dbodo
Heieb
Dbodo
Sch sp
Heieb
Dic/Ris
Equsp
Dbodo
DidE_gu_ _
Hei eib
Equsp
Heieb
_Hej/Dic _
Sph pon
Dic/Ris
Sch sp
. Equsp _
Db/Hei
Equ sp
Dbzub
Db2spp
_Hei eb_ _
Dbzub
Dbodo
Dbqa
Hei/Dic
Dbzub
Equsp
Db3spp
Dbeb
Db2spp
Dbodo
Db dub
_Dicodq_ _
Db dub
Dbcra
Db 2spp
Hei/Db
Dbdub
Dic/Sph
Dbeb
Equ sp
Dbdub
_75 _ 6
1
2 51 3
1 38 14 7 47
18 - - 26 27
Tab. 6b. Molteno gymnosperms, reproductive genera (abundance & frequency)
see p. 9 for notes
Sampling
12
Gj/tRELITZIA 15 (2003)
2. FREQUENCY & ABUNDANCE
Differentiating frequency & abundance
The measures of frequency and abundance are crucial in our
endeavour to uncover diversity. Our usage of the terms is quite
specific within the context of this work.
Frequency. The measure of frequency of a taxon within the
Molteno Fm. is the number of taphocoenoses (TCs or assem-
blages), out of the total 100 sampled, in which it has been found.
Through the Gondwana Triassic it is the number of productive
degree squares (subregions), out of the total 85 sampled, in which
it has been found.
Abundance. The abundance of a taxon is a measure of the
absolute or relative number of individuals recorded or collected
from an assemblage, formation, region or continent.
In Gondwana Triassic studies, precise abundance data are
available only for the Molteno Fm. For other Gondwana conti-
nents, where clear counts of collected individuals are rarely given
in the literature, the figures recorded below (Tabs 7a,b) and in the
hypodigm tables scattered through the systematic section, are the
number of illustrated individuals.
Recording abundance
Fruit. As recorded later (Tabs 9a,b-I2), reproductive organs are
preserved very much less abundantly than foliage.
Absolute abundance (curated individuals)'. The number of individual
specimens of fruit counted in the curated collection.
Relative abundance (counts per man-hour or day)'. This is a measure
of the rate of appearance of taxa within particular TCs. It is recorded
as the number of individuals encountered— while cleaving fossilifer-
ous slabs on site or in the lab— within an hour or day ( 10 hours). While
recognising that collectors work at different rates and with varying
acuity, these figures are readily attained and highly instructive.
Foliage
Absolute abundance : The tally of curated individuals of a taxon
where it accounts for < 1% of the total vegetative fraction of the
assemblage (TC).
Relative abundance'. The percentage occurrence (estimated at site)
of a taxon where it accounts for 1 % or more of the vegetative frac-
tion of the TC.
Ovulate genera
CLASS
ORDER
Genus
Individual counts
| SAm
Molt, t
It*
other if
"O
c
Aus
Ant
PINOPSIDA (Coniferopsida)
DORDRECHTITALES
Dordrechtites
-
413
-
-
4
-
VOLTZIALES
Voltziopsis*
-
-
1
-
6
-
Telemachus
3
311
-
-
4
5
PINALES (Coniferales)
Rissikistrobus
-
85
-
-
1
-
Gypsistrobus
-
5
-
-
-
-
Avistrobus
-
1
-
-
-
-
Parasciadopitys*
-
-
-
-
-
2
GINKGOOPSIDA
PELTASPERMALES
Peltaspermum
2
257
-
-
3
-
MATATIELLIALES
Matatiella
-
17
-
-
24
-
GINKGOALES
Karkenia*
-
-
-
-
1
-
Avatia
-
114
-
-
-
-
HAWSHAVWIALES
Hamshawvia
1
24
-
-
2
-
UMKOMASIALES
Umkomasia
9
503
-
-
61
1
Fanerotheca
13
247
-
-
-
-
PETRIELLALES
Petriellaea*
-
-
-
-
-
HT
Kannaskoppia
-
50
-
-
-
-
ORDER indet.
Cetifructus
-
2
-
-
-
-
CLASSES indet.
ALEXIALES
Alexia
-
6
-
-
-
-
HLATIMBIALES
Hlatimbia
-
2
-
-
-
-
ORDER indet.
Hystricia
-
1
-
-
-
-
BENNETTITOPSIDA
FREDLINDIALES
Fredlindia
-
16
-
-
5
-
PENTOXYLALES
Lindtheca
GNETOPSIDA
NATALIGMALES
Nataliqma
-
4
-
-
-
-
FRAXINOPSIALES
Fraxinopsis
40
306
-
-
3
-
Total individuals (approx.)
68
2378
1
-
114
19
Total genera
6
20
1
-
11
4
Microsporangiate genera
CLASS
ORDER
Genus
Individual counts
E
<
w
<-SA
o
5
f->
0)
£
o
■O
c
</)
3
<
c
<
PINOPSIDA (Coniferopsida)
VOLTZIALES
Fredianthus
-
2
-
-
-
-
Lutanthus
-
5
-
-
-
-
Odyssianthus
-
2
-
-
-
-
PINALES (Coniferales)
Rissikianthus
-
79
-
-
-
-
ORDER indet.
Flelvetianthus
-
6
-
-
-
-
CYCADOPSIDA
CYCADALES
Androstrobus
-
2
-
-
-
-
GINKGOOPSIDA
PELTASPERMALES
Antevsia
-
32
-
-
-
-
MATATIELLALES
Switzianthus
-
54
-
-
-
-
GINKGOALES
Eosteria
-
27
-
-
-
-
HAMSHAVWIALES
Stachyopitvs
4
539
-
-
3
-
UMKOMASIALES
Pteruchus
5
425
-
i
14
1
PETRIELLALES
Kannaskoppianthus
-
92
-
-
?1
-
BENNETTITOPSIDA
FREDLINDIALES
Cycadolepis
-
14
-
-
-
-
Weltrichia
-
3
-
-
-
-
Leguminanthus
-
5
-
-
-
-
Total individuals
9
1287
-
i
17
1
Total genera
2
15
-
i
3
1
Tabs 7a, b. Gondwana Triassic (GT) gymnosperms, ovulate &
microsporangiate genera
Molteno counts', tally of curated individuals
Non-Molteno counts', includes all illustrated individuals in the Gondwana
Triassic literature (based on hypodigms published here); clear counts
of collected individuals are rarely given in the literature
Generic list: accounts for all GT genera recognised here;
* the 4 ovulate genera Voltziopsis. Parasciadopitys, Karkenia and
Petriellaea, unknown in the Molteno, are marked by an asterisk
SAm — South America Ind— India Ant — Antarctica
SAf — Southern Africa Aus — Australasia
Frequency & abundance
^/tRELITZIA 15 (2003)
13
Abundance
in a single TC
Percent- | Indivs per man-hours cleaving where < 1% | Indivs per man-days
age ' metric heirarchy | range 1 of cleaving
monodominant
co-dominant
abundant
common
sparse
1 i ~ i
70-ioo%i ; - |
20-69% ! - 1 !
6-19% | - ! !
3-5% ; - ; - |
1 !_
rare
very rare
extremely rare
vanishingly rare
infinitely rare
i >1 per 1 man-hour 1 >1 per <5 man-hours j >1 per < V2 man-day
- 1 ca 1 ” 10 man-hours 1 1 ” 5-49 ” 1 1 ” 1-4 man-days
! ca 1 " 100 ” | 1 ” 50-499 ” ! 1 ” 5-49
| ca 1 " 1000 ” | 1 ” 500—4999 ” ! 1 " 50-499
| ca 1 " 10000 ” | 1 in over 5000 " | 1 in over 500
1 1 !
Abundance
in a single TC
Examples
Genus TC
% in
TC
1 1
1 Rate in man-hours [ Rate in man-days
1 (10 man-hours cleaving = 1 man-day)
monodominant
Heidiphyllum
in Aas 311
Hei elo 1
99%
1
1
1
1
co-dominant
Dicroidium
” Lit 111
Dic/Hei |
50%
1
-
1
abundant
Halleyoctenis
" Kon 223
Die odo |
15%
1
-
common
Sphenobaiera
” Kon 111
5%
1
-
-
sparse
Lepidopteris
” Lit 111
Dic/Hei !
1%
1
-
1 -
rare
Stachyopitys
" Kon 222
Die odo |
-
| 36 indivs in
40 man-hours
| 9 indivs in 1 man-day
very rare
Fredlindia
-
| 3 indivs in
40
| ca 1 " ” 1 "
extremely rare
Fredlindia
” Aas 411
Dic/Sph '
-
' 11 indivs in
512
1 ca 1 " " 5 ” days
vanishingly rare
Hystricia
1
-
1 1 indivs in
512
1 ca 1 ” ” 51
infinitely rare
-
1
1
-
1
1
-
1
Tabs. 8a, b. Abundance scale, foliage and fertile organs
Abundance: 10 categories are applied; the top five (>1%) reflecting relative abundance and the lower five
(<1%) absolute abundance.
Percentage: This is an estimate made in the field (less often the laboratory) for taxa yielding over 1 % of the total
assemblage. In those few instances where fertile taxa occur commonly to abundantly on certain bed-
ding plains (e.g. Dordrechtites scales), the actual tally of curated individuals is nevertheless given on the
various relevant tables.
Examples: Since foliage occurs very much more abundantly than fruit, the former are used to illustrate the
upper half of the scale and the latter the lower half. Both categories of organ may, however, fall any-
where within the scheme.
Individuals per man-hours cleaving: as noted, the lower half of the scale applies primarily to fruit, less often
foliage. For less thoroughly sampled TCs, all or nearly all individuals are collected, curated and counted. As
the intensity of sampling any particular TC increases, so the selectivity for better preserved specimens
increases. This process for retention is progressive and commences at ca 20 individuals (e.g.
Fraxinopsis at Kap 111). Where higher figures are indicated (e.g. >50), the retention rate is ca 50% (e.g.
Pteruchus at Mat 111) and for very common species (e.g. Dordrechtites scales at Aas 411) the retention
figure may drop as low as 10%.
Ovulate
genera
TCs
Indivs
Ref. TC
1
Fanerotheca
27
247
Bir 111
2
Umkomasia
22
503
Mat 111
3
Telemachus
18
311
Tel 111
4
Fraxinopsis
18
306
Bir 111
5
Dordrechtites
17
413
Lut 311
6
Peltaspermum
17
257
Bir 111
7
Rissikistrobus
7
85
Umk 111
8
Avatia
6
114
Bir 111
9
Hamshawvia
4
24
Umk 111
10
Matatielia
4
17
Mat 111
11
Fredlindia
3
16
Aas 411
12
Kannaskoppia
1 50
Kan 111
13
Lindtheca
1 16
Aas 411
14
Alexia
1 6
Umk 111
15
Gypsistrobus
1 5
Aas 411
16
Nataligma
1 4
Umk 111
17
Hlatimbia
1 2
Hla 213
18
Cetifructus
1 2
Umk 111
19
Hystricia
1 1
Aas 411
20
Avistrobus
1 1
Bir 111
Totals
2378
Microsporangiate
genera
O
1-
Indivs
Ref. TC
1
Stachyopitys
27
539
Bir 111
2
Pteruchus
22
425
Umk 111
3
Kannaskoppianthus
12
92
Lut 311
4
Rissikianthus
5
79
Pen 321
5
Antevsia
5
32
Maz 211
6
Switzianthus
4
54
Lit 111
7
Eosteria
4
27
Aas 311
8
Cycadolepis
3
14
Kon 222
9
Lutanthus
3
5
Lut 311
10
Weltrichia
2
3
Kon 222
11
Androstrobus
2
2
Pen 321
12
Helvetianthus
1 6
Lit 111
13
Leguminanthus
1 5
Kon 222
14
Odyssianthus
1 2
Tel 111
15
Fredianthus
1 2
Aas 411
Totals
1287
Tabs 9a, b. Molteno fertile genera, frequency and abundance
Genera: arranged according to decreasing frequency, then decreasing
abundance, then man-hours (see Tab. 11)
Taphocoenoses (TCs): number of TCs in which genus is known
Individuals (indivs): total tally of specimens from all TCs
Reference taphocoenosis (RTC): that TC yielding the best sampled
palaeodeme(s) for the genus
Total individuals: 9 ca 2378; cf ca 1287
Frequency & abundance
14
^TRELITZIA 15 (2003)
Relative abundance of the rarest genera Microsporangiate genera. The four least frequent and abundant
, , , microsporangiate genera in the Molteno, Leguminanthus (1 per 10
0vulate genera. The nine least frequent and abundant ovulate man_hours), Odyssianthus (1 per 45 man-hours); Helvetianthus (1
genera in the Molteno-each occurring in only one TC-vary con- per 10Q man_hours), and Fredianthus ( i per 250 man-hours), range
siderably in absolute (1-50 individuals) and relative abundance on our abundance scale from <very rare- t0 ‘extremdy rarey
(Tabs 9a, 10). Following our scale of abundance (Tab. 8), The rarest of all described microsporangiate species in the
Kannaskoppia is ‘rare', appearing at a rate of two individuals per Molteno is Lutanthus robustus, with one individual from Aas 4 1 !
man-hour; Hlatimbia and Lind theca are 'very rare: Alexia , ( j pCr 5 1 2 man-hours). As two other species of Lutanthus occur at
Natal, gma, Gypsistrobus and Cetifructus. each appearing at a rate tWQ further TCs> lt is not among the least frequent 0f genera,
of one individual within the range 5CM-99 man-hours, are all
‘ extremely rare' , while Hystricia and Avistrobus are the rarest of Foliage genera. Of the 27 (+ 1 unnamed) gymnospermous foliage
all recorded Molteno ovulate genera, each with only one fertile genera recognised in the Molteno, fully one quarter (6 plus 1
head from one TC in over 500 man-hours cleaving. unnamed) are known from only a single TC. These scarcest foliage
Statistical projections suggest that many more genera, still elements range in relative abundance from very rare through
rarer than Hystricia and Avistrobus. are preserved in the Molteno, extremely rare to vanishingly rare, following much the same pattern
but are yet to be observed (Anderson et al. 1 996). For the most elu- as the reproductive genera. Saportaea, exclusive to Little Switzerland
sive of these, the category of ‘ infinitely rare' , i.e. one individual in (Lit 111) and with only a single specimen found in 550 man-hours
over 5 000 man-hours, is available. cleaving, is the rarest of all foliage genera in the collection.
Ovulate genus
TC
Indivs
Sampling intensity
Relative abundance
Abundance scale
Kannaskoppia
Kan 111 Ast spA
50
30 man-hours
2 per 1 man-hour
rare
Hlatimbia
Hla 213 Die elo
2
60 ”
1 per 30 man-hours
very rare
Lindtheca
Aas 411 Dic/Sph
16
512 ”
1 per 40
Alexia
Umk 111 Die 2spp
6
400 "
1 per 66
extremely rare
Nataligma
Umk 111 Die 2spp
4
400 ”
1 per 100 ’’
” ”
Gypsistrobus
Aas 411 Dic/Sph
5
512 ”
1 per 102 ”
Cetifructus
Umk 111 Die 2spp
2
400 ”
1 per 200 ”
” ”
Hystricia
Aas 411 Dic/Sph
1
512 ”
1 per 512 ’’
vanishingly rare
Avistrobus
Bir 111 Sph 2spp
1
550 ”
1 per 550 ”
Microspor. genus
TC
Indivs
Sampling intensity
Relative abundance
Abundance scale
Leguminanthus
Kon 222 Die odo
5
40 man-hours
1 per 10 man-hours
very rare
Odyssianthus
Tel 111 Hei elo
2
90 ”
1 per 45
Helvetianthus
Lit 111 Dic/Hei
6
550 ”
1 per 100 ”
extremely rare
Fredianthus
Aas 411 Dic/Sph
2
512 ”
1 per 250 ”
Foliage genus
TC
Indivs
Sampling intensity
Relative abundance
Abundance scale
Jungites
Lit 111 Dic/Hei
18
550 man-hours
1 per 30 man-hours
very rare
Graciliglossa
Umk 111 Die 2spp
13
400 ”
1 per 30
unnamed gnetalean
7
400 ”
1 per 60
extremely rare
Cetiglossa
3
400 ”
1 per 1 30 ”
Pagiophyllum
Aas 411 Dic/Sph
2
512 ”
1 per 250 ”
Scytophyllum
Umk 111 Die 2 spp
1
400 ”
1 per 400 ”
Saportaea
Lit 111 Dic/Hei
1
550 ”
1 per 500 ”
vanishingly rare
Tab. 10. Relative abundance of the rarest Molteno genera
Genera listed: only those genera with a frequency of 1 (occurring in a single TC)
are included
Ovulate & microsporangiate genera: for frequency and abundance data
see Tabs 6b & 9; and for man-hours cleaving Tab. 1.
Foliage genera: for frequency and abundance data see Tab. 6a; and for man-
hours cleaving Tab. 1 . (See Tab. 1 2 for further details on the unnamed
gnetalean.)
Frequency & abundance
^/trELITZIA 15 (2003)
15
assemblages
(taphocoenoses)
Sampling
C .C O)
ii 15
Ovulate genera
I 5 5
« £
5 ! '5 1 1
<TJ Q Q. f- o
E g ® Q) Q.
| I 'j 5 3 : » 1 i I % I
E a J o o>-I2s<v<52’(!]
U.Q Q.CC'tlSu.ic
•E -Si
o 2 i: o a:
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20
Microsporangiate genera
■Q -C C £
<U £ *r
5!
1 23456789 10 11
Aas 411
Dic/Sph
512
2176
Umk 111
Die 2spp
400
3592
Lit 111
Dic/Hei
550
2173
Bir 111
Sph 2spp
550
1927
Kap 111
Die spp
65
1310
Lut 311
Hei elo
50
589
Maz 211
Hei/Dic
85
1087
Mat 111
Die dub
65
1082
Kon 222
Die odo
40
449
San 111
Die era
30
303
Bir 311
Hei/Sph
2
133
Hla 213
Die elo
60
902
Ela 111
Die odo
10
235
Cyp 111
Die era
100
362
Tel 111
Hei elo
90
581
Pen 411
70
204
" 311
n
35
155
Kle 111
Hei/Dic
9
288
Bir 211
Sph 2spp
7
184
Tin 121
"
5
33
Wal 111
Die odo
50
339
Pen 321
Dic/Ris
35
408
Kon 111
Die odo
34
285
Maz 111
Die era
30
462
Kan 111
Ast spA
30
365
Gre 111
Equ sp
25
213
Nuw 111
Die zub
21
138
Kra 1 1 1
Die odo
20
380
Hla 212
Die 3spp
11
217
Boe 112
Die cor
6
179
Mol 111
Sph pon
6
95
Kon 111
Hei elo
4
55
Hla 211
Die 3spp
4
157
Aas 311
Hei elo
140
528
” 111
" "
40
291
” 211
” "
35
154
Win 111
Hei elo
20
64
Kan 112
” "
15
145
Kle 111
n
15
15
Ask 111
Equ sp
14
366
Kra 311
Die odo
13
159
Gol 111
Die dub
13
108
Mor 111
Die odo
12
214
Gre 121
Hei elo
10
141
Korn 111
Sph/Dic
10
168
Qua 111
Die odo
8
140
Boe 111
Dic/Hei
8
98
” 111
Lep sto
8
77
Nuw 211
Die 2spp
6
119
Qac 111
Hei/Dic
4
112
Ela 112
Dic/Hei
4
41
Aas 611
Hei elo
3
47
Lut 411
Hei/Dic
2
60
" 111
2
23
" 511
Hei elo
2
26
Nav 111
Die odo
2
60
Vin 111
Die odo
10
166
Kle 111
Equ sp
4
35
57 TCs
Total TCs
44 1 00 88 20 38 1 4 1 11 - 16 - 5 - - -
J4 _10_17_45_ _7_ 6_ 4_ 2_
10 50 11 8 - 2
60 3 21 15 -
J9 1_38__-_ 8_ 7
4 36 9 - V
113 --- -
35 11 - 25 -
19 __ _- _16_ -
26 27 - - 18 *
31 84 3 - -
35 13 - - 2
7 5 - - 1
11-..
26 -
_6 _50
63 40
67 - 66 -
1 3 - -
7 - 3 1
46
75 12
2 20
_1_ _1_
2 5
6443111111 111
- 27 22 12 5 5 4 4 3 3 2 2 1 1
Tab. 11. Molteno ovulate & microsporangiate genera, sampling intensity & biodiversity
Taphocoenoses: the 57 TCs are arranged according to diversity of ovulate genera, then to sampling
intensity (man-hours cleaving)
Sampling: two measures of sampling intensity for each TC are recorded (see text for explanation)
Genera: arranged in order of decreasing frequency, then decreasing abundance (see Tabs 9a, b)
Matrix: the absolute abundance (individuals) perTC is recorded
Long-tailed frequency & diversity curves: note the long-tailed curves for both generic frequency in the Molteno
and generic diversity per TC — which point to many ( preserved) genera still awaiting discovery
Frequency & abundance
16
^TRELITZIA 15 (2003)
3. AFFILIATED ORGANS
Reassembling the whole plant
A comprehensive strategy towards seeking out affiliations
between dispersed organs in a fossil flora is essential to gaining an
insight into the true (natural) diversity ( observed , preserved and
existed) of that flora (Anderson et al. 1996). It is inevitable that the
reliability of proposed links between organs will vary consider-
ably—from marginal likelihood of affiliation (Grade 1) to certain-
ty of affiliation through organic attachment (Grade 5). The relia-
bility of each foliage, female or male affiliation documented here
is indicated according to the following system.
Criteria for affiliations (elaborated after And. & And. 1985, p. 85)
Judgements concerning affiliations are based on an array of
observations. Reliability will depend on the following criteria.
(The abbreviations used throughout this volume are given in
brackets.)
1. Organic attachment (Org. att.) — Organs that are found in
direct organic connection constitute the only irrefutable case for
conspecific status.
2. Cuticle correspondence (Cut. cor.) — It is reasonably estab-
lished that the cuticles of different organs of the same species dis-
play like characteristics.
3. Morphological correspondence (Mor. cor.)— In certain instances,
diagnostic macroscopic features such as ornamentation, blistering
and texture are seen in conspecific organs.
4. Kindred reinforcement (Kin. rein.)— Well authenticated organ
affiliations for other genera in the family or order offer a secure
foundation for proposing linkage.
5. Mutual occurrence, presence or absence (Mut. occ.)— Where
different dispersed organs occur in the same assemblage, the pos-
sibility exists that they derive from the same parent species. The
likelihood will increase with:
• coupling frequency— the number of assemblages in which the
mutual occurrence is repeated;
• mutual abundance— the mutual dominance or rarity of the organs
in question;
• process of elimination — the preoccupation of organs in other
established affiliations;
• bedding-plane bonds— the extent to which the organs are con-
fined to particular bedding planes;
• assemblage paucity— the lowering of diversity levels;
• assemblage autochthony — the degree to which the assemblage
represents a single, local plant association.
Reliability grades (after And. & And. 1985, p. 85)
The evidence for linking organs ranges from marginal to cer-
tain. At the lower end of the range the evidence will be slim, yet
suggestive, or alternate options might be more or less equally likely,
while at the upper end of the range clear organic attachment certi-
fies linkage.
Grade 1, marginal -
Grade 2, poor
Grade 3, fair -
Grade 4, good -
Grade 5, certain -
Marginal likelihood of affiliation:
mutual occurrence (weak).
Most feasible affiliation (alternatives competitive):
mutual occurrence (unclear).
Probable affiliation (alternatives weak):
mutual occurrence (fairly clear),
little other supportive data.
Virtually exclusive likelihood of affiliation:
mutual occurrence (particularly clear),
cuticle correspondence and/or kindred reinforcement
and/or possible organic attachment.
Certain affiliation:
organic attachment undoubted.
Organic attachment is known for only five gymnospermous genera
(generic pairs) in the Gondwana Triassic, four from the Molteno and
one from Australia.
1 . Voltziopsis— NSW, L. Triassic (Townrow 1967b, pi. le)
2. Kannaskoppia/Kannaskoppifolia— Molteno (this vol.)
3. Kannaskoppianthus/Kannaskoppifolia— Molteno (this vol.)
4. Hamshawvia/Sphenobaiera—Moheno (this vol.)
5. Stachyopitys/Sphenobaiera— Molteno (this vol.)
Lhnkomasia unirama (Axsmith et al. 2000) falls into affiliation Grade
4 as there are two separate shoots, one bearing the fruit and another the
leaves.
Organic attachment, Gondwana Triassic
Extreme rarity of (foliage-fruit) organic attachment
Organic attachment is an exceptionally rare phenomenon in the
plant fossil record. Only four instances of mutual attachment,
involving two whole-plant genera, occur in our Molteno collec-
tions that approach 30 000 catalogued slabs from 100 TCs. In the
first instance, several specimens from a restricted pocket of silty
sediment (at Kan 1 1 1 Ast spA) occur, while in the latter three cases
(at three different TCs) only single specimens have been found.
Just two whole-plant genera are involved.
Kannaskoppia/Kannaskoppifolia! Kannaskoppianthus
Kan 111: Kannaskoppia/Kannaskoppifolia ( pi . 104)
This taxon provides the finest example, from amongst the wide
diversity of gymnospermous plants found in the formation, of
specimens that clearly show organic attachment between foliage
and female fruit. Several specimens of slender stems from Kan 1 1 1
Ast spA bear short shoots with both leaves and female strobili
(Grade 5 affiliation).
Kom 111: Kannaskoppianthus/Kannaskoppifolia (pi. 108, 109)
One further TC (Kom 1 11 Sph/Dic) has yielded a single speci-
men consisting of four Kannaskoppifolia leaves and two undoubted
strobili of Kannaskoppianthus attached to a shoot. An additional
28 specimens of individual leaves of the same species, but different
from those from Kan 111, occur in the assemblage.
Hamshawvia/SplienobaieralStachyopitys
Aas 411: Hamshawvia/Sphenobaiera (pi. 71)
From Aas 4 1 1 Dic/Sph, one specimen shows a cluster of Spheno-
baiera leaves and a small bifurcate female strobilus attached to the
terminal portion of a shoot. The incomplete strobilus is compara-
ble to Hamshawvia but clearly smaller and is most likely either an
immature or undeveloped specimen of that genus. In view of this
uncertainty, we rate the affiliation Grade 4/5 rather than 5.
Maz 111: Stachyopitys/Sphenobaiera (pi. 8 1 )
A single specimen from Maz 1 1 1 Die era shows a Stachyopitys
strobilus (male) and an incomplete Sphenobaierct leaf attached to a
bulbous base (Grade 5 affiliation).
There exists supporting evidence from Australia and Eurasia
(pp. 213, 218) for the mutual occurrence or attachment of Spheno-
baiera and its reproductive counterparts.
Affiliated organs
c9 TRELITZIA 15 (2003)
17
Relative occurrence of different organs
Paucity of male versus female strobili
It is a conspicuous feature seen in the gymnosperm component
of many fossil floras that the male strobili are less diverse, less fre-
quent and less abundant than the female strobili. This is true also
of the Molteno flora, but not so markedly as we initially thought.
The further our studies of the reproductive structures have pro-
gressed, the more prominent has become the male presence. As
indicated in the text table below, the females outnumber the males
in all respects, but the actual discrepancy varies significantly. As
one works down through the taxonomic ranks, from class to genus,
the deviation between female and male in observed diversity
diminishes steadily and markedly (from 2:1 down to 1.3:1). At
species level it increases again slightly to 1.5:1. Females outnum-
ber the males in frequency (at a ratio of 1.6:1) and in abundance
(2: 1 ). The shape of the frequency curves (distributions) for ovulate
and microsporangiate strobili is very similar (Tab. 11) in that both
decline rapidly from occurrences in ca 25% of TCs to a long tail of
rare to single appearances.
We note two further general examples reaffirming this relative
paucity of male cones. The first is in the order Voltziales, promi-
nent globally from the U. Permian to M. Jurassic, and the second
in the order Glossopteridales, dominant throughout the Gondwana
Permian. In the Voltziales only three genera of male strobili to 15
female genera are known (And. & And. 1989. p. 422). In the
Glossopteridales the proportion of described male to female gen-
era is one to ten (And. & And. 1985, p. 107).
Paucity of strobili versus foliage
The vegetative component in compression-impression floras
such as those preserved in the Molteno far outstrips in all measures
the reproductive component. The discrepancy increases rapidly
from class (foliage:female = 8:8 = 1:1) through to species
(foliage:female = 113:51 = 2.2:1). This will be a reflection partly
of straight biomass (frequency and abundance) of the original liv-
ing material (biocoenosis) and partly due to differential taphonom-
ic filtering (reflected in the taphocoenosis). Foliage palaeodemes
outnumber ovulate palaeodemes 3:1 (440:152), while foliage indi-
viduals outnumber ovulate individuals (excluding dispersed seeds
or scales) at least in the order of 100:1 (250 000:2 378).
Foliage/fruit discrepancy in species diversity
We consider here (following a particular case study) the appar-
ent discrepancy between foliage, female and male species diversi-
ty. The pattern for species of Molteno Ginkgoopsida— a clear
reversal of that seen in the Pinopsida— is for greater diversity in the
foliage than in the female or male fruit. The whole-plant
Peltaspermum! Lepidopteris presents an exception to this rule: for
Peltaspermum we define five species, for Lepidopteris only two
species, and for Antevsia one species. The male, Antevsia, occurs
particularly infrequently and rarely, and the affiliation with
Lepidopteris is less sure (Grade 3).
What is the reason for this apparent Peltaspermum-Lepidopteris
diversity anomaly? Is it simply a reflection of imperfect taxonom-
ic decisions reflected in our monograph (And. & And. 1989) on the
gymnospermous foliage of the Molteno a decade ago, when our
Peltaspermum collections were significantly less complete? We
witness here that the morphologically simpler P. monodiscum
occurs far more widely in the Molteno than the more complex P.
tridiscum , P. turbanatum and P. quindiscum, which appear to have
been restricted to different specific habitats. In referring back to the
foliage of the relevant localities (TCs), the question is whether we
can now identify further morphological differences suggesting
greater species-level diversity. Does the cuticle contribute to
resolving the issue? These issues are partly addressed below.
Little Switzerland (Lit 111) & Umkomaas Valley (Umk 111): The
excellently preserved cuticle from these two TCs clearly supports
the identification of two Lepidopteris species, L. africana and L.
stormbergensis. From the patterns of occurrence of foliage and
fruit at Lit 111 it is most readily interpreted that P. monodiscum
might affiliate with L. africana and P. quindiscum with L. storm-
bergensis, while at Umk 111, L. thomasii might affiliate with L.
africana and P. quindiscum with L. stormbergensis.
Birds River (Bir 111 ) & Aasvoelberg (Aas 411): These two lake-
deposit TCs both yield the two Peltaspermum species P. monodis-
cum and P. turbanatum. The Lepidopteris palaeodemes from each
are well-represented and diverse and may well include more than
one natural species (all are included as L. stormbergensis at Bir 1 1 1
in And. & And. 1989).
Konings Kroon (Ron 111. Ron 222) & Peninsula (Pen 321): The
Lepidopteris palaeodemes from Kon 111 (pis 41, 42) and Kon 222
(pi. 43) are extensively illustrated in And. & And. ( 1989). The lat-
ter, yielding the reference palaeodeme (24 individuals) of P. tri-
discum, includes foliage only of the general L. africana range-
suggesting that this ‘species’ affiliates with P. tridiscum.
Molteno gymnosperms
T
1
1
Whole-
9 : cf
plant
Prominence
Ranks
foliage
female
i male
deviation
taxa
Diversity (observed)
classes
8
8
4
2 : 1
10
orders
17
18
i ii
1,6 : 1
23
families
24
18
i 13
1,4 : 1
32
genera
27
20
i is
1,3 : 1
38
species
113
51
! 35
1,5 : 1
143
Frequency
palaeodemes
440
152
1 94
1,6 : 1
-
Abundance
individuals
ca 250,000
2378
! 1288
2 : 1
-
Molteno gymnosperms, diversity, frequency & abundance of the three plant organs
Taxonomic ranks: from class to individual
Whole-plant taxa: for classes, orders & families refer to Tab. 30, pp. 54, 55;
for genera & species refer to Tab. 15, p. 21 .
Affiliated organs
18
d^TRELITZIA 15 (2003)
Ovulate
Foliage
Microsporangiate
WPG
genera
(/) 0
in -2 i
O ? t!
I— .E too
genera
TCs
indivs
genera
TCs
indivs
affil.
grade
1
Kannaskoppia
1 50 5
Kann. (folia)
1
25
Kann. (anthus)
12 92 5
2
Hamshawvia
4 24 14/5
Sphenobaiera
43
Stachyopitys
27 539 5
3
Umkomasia
22 503 ; 4
Dicroidium
75
Pteruchus
22 425 4
4
Telemachus
18 '311 4
Heidiphyllum
62
Odyssianthus
2 1 2 < 4
5
Rissikistrobus
8 85 ! 4
Rissikia
21 '
Rissikianthus
5 79 4
6
Peltaspermum
17 1 257 4
Lepidopteris
30
Antevsia
5 32 3
7
Fredlindia
3 i6 ; 3
Halleyoctenis
10
Cycadolepis 1
3 14 3
”
1 1
1
Weltrichia
2 ; 3 ; 3
8
Avatia
6 114 2
Ginkgoites
19 : 250
Eosteria
4 : 27 ; 3
9
Fraxinopsis
18 '306 4
Yabeiella
29
-
- 1 - 1 .
10
Lindtheca
1 16 3
Taeniopteris
38
-
11
Nataligma
1 ! 4 3
Gontriglossa
8 . 100
-
i i
12
Fanerotheca
26 247 2
Dicroidium
? | ?
-
i i
13
Matatiella
4 17 ! 2
Kurtziana
13 150
-
_ i _ i
14
Hlatimbia
1 2 1 2
Batiopteris
o
->J
o
- ! - ! -
15
-
- 1 - 1
Dejerseya
5 1 200
Switzianthus
4 54 : 2
16
-
Pseudoctenis
21 ' 250
Androstrobus
2 ! 2 ! 2
17
-
1 I
Linguifolium
9 75
-
1 1
18
-
Moltenia
5 55
-
_ 1 _ 1
19
-
- ! . i .
Clariphyllum
3 ' 51
-
. ! . ! .
20
-
Jeanjacquesia
3 ' 8
-
- i - i
21
-
1 i
Paraginkgo
2 43
-
22
-
I i
Ctenis
2 : 3
-
I I
23
-
. i . i .
Jungites
1 1 18
-
_ l _ i
24
-
. ! - i -
Graciliglossa
1 13
-
- ! . ! .
25
-
- i - i -
Cetiglossa
1 3
-
- ! -\ -
26
-
Pagiophyllum
1 ; 2
-
. 1 _ 1
27
-
I i
Scytophyllum
1 1
-
28
-
Saportaea
1 1
-
29
Dordrechtites
17 413 ' -
-
. 1
-
-i - i -
30
Alexia
1 6 ' -
-
- !
-
- 1 - i
31
Gypsistrobus
1 5 ! -
-
-
■ i " i
32
Cetifructus
1 I 2 1 -
-
I
-
i i
33
Hystricia
11
-
I
-
_i . i
34
Avistrobus
1 1
-
- !
-
- i - i
35
-
- | - 1 -
-
- 1
Lutanthus
3 ! 5 -
36
-
-
Helvetianthus
1 6 i -
37
-
I 1
-
1
Leguminanthus
i ; 5 1 -
38
-
-
1
1
Fredianthus
i ; 2 : -
20 genera
1 1
1 »
27 genera
1
1
15 genera
l i
1 I
Tab. 12. Affiliated organs in the Molteno, statistics
Frequency: number of TCs of 100 sampled in Molteno
Abundance: tally of individuals in curated collection;
generally rounded off where >50 (in case of foliage)
= numerous (far >1000 leaves)
= many (>500 leaves)
Affiliation: reliability grade 1 to 5
Whole-plant genera (WPG): in order of affiliation grade, then frequency
(TCs), abundance (indivs), & relative abundance (collecting
intensity) of female fruit
Organ-genera: in order of frequency (TCs), then abundance (indivs)
Unnamed gnetalean foliage genus: this genus [see And. & And. 1983,
Ginkgophytopsis spD from Umk III, pi. 10(6)] remains unde-
scribed in the present work; and, with 7 indivs in a single TC,
should fall in this table between WPGs 24 & 25 (see Tab. 10
for further details).
Biodiversity implications
The scarcity of established affiliations (Tab. 12)
• Of the 38 gymnospermous whole-plant genera recognised in
the Molteno, only eight are known from all three organs —
female, foliage and male.
• For a further eight whole-plant genera, either female/foliage (6
cases) or foliage/male (2 cases) affiliations are established.
• A total of only 16 of 38 whole-plant genera are thus represent-
ed by more than one of the three organs (i.e. 16 multi-organ
genera are known).
• Six female genera, four male genera and 12 foliage genera
remain without any hint of their affiliated organs.
From observed to preserved diversity
The implications concerning biodiversity that flow from the
rarity of reasonably established affiliations are of particular signif-
icance in this study. If less than half of all observed Molteno
whole-plant genera are known from two or three organ-genera ,
and over half are based on only a single organ, a long tail of whole-
plant genera not represented at all can be projected. Considering
the high diversity of preserved taxa statistically extrapolated from
frequency distributions of observed taxa (p. 25), the extent of this
tail is, indeed, shown to be long.
Affiliated organs
TRELITZIA 15 (2003)
19
DIVISION
CLASS
ORDER
Genus
Ovulate
affiliation
grade
Foliage
affiliation
grade
Microsporangiate
PINOPHYTA (gymnosperms)
PINOPSIDA (Coniferopsida)
DORDRECHTITALES
Dordrechtites
—
—
—
-
VOLTZIALES
—
—
—
—
Fredianthus
-
—
-
—
Lutanthus
Tetemachus
4
Heidiphyllum
4
Odyssianthus
-
—
Clariphyllum
-
PINALES (Coniferales)
Rissikistrobus
4
Rissikia
4
Rissikianthus
—
—
Pagiophyllum
—
-
ORDERS indet. (2 total)
Gypsistrobus
-
—
—
-
Avistrobus
—
-
—
-
—
—
—
—
Helvetianthus
CYCADOPSIDA
CYCADALES
-
-
Pseudoctenis
2
Androstrobus
-
—
Jeanjacquesia
—
-
-
—
Ctenis
—
-
—
—
Moltenia
—
-
GINKGOOPSIDA
PELTASPERMALES
Peltaspermum
4
Lepidopteris
3
Antevsia
—
—
Scytophyllum
—
-
MATATIELLALES
Matatiella
2
Kurtziana
—
-
-
-
Dejerseya
3
Switzianthus
GINKGOALES
Avatia
2
Ginkgoites
3
Eosteria
-
—
Paraginkgo
—
-
HAMSHAWVIALES
Hamshawvia
4/5
Sphenobaiera
5
Stachyopitys
UMKOMASIALES
Umkomasia
4
Dicroidium
4
Pteruchus
Fanerotheca
2
DicroidiumJ
-
PETRIELLALES
Kannaskoppia
ORDER indet
5
Kannaskoppifolia
5
Kannaskoppianthus
Cetifructus
—
—
—
—
CLASSES indet. (5 total)
ALEXIALES
Alexia
—
—
—
-
HLATIMBIALES
Hlatimbia
2
Batiopteris
-
-
ORDERS indet. (3 total)
Hystricia
-
-
-
-
-
—
Saportaea
—
-
—
—
Linquifolium
—
-
BENNETTITOPSIDA
FREDLINDIALES
Fredlindia
3
Halleyoctenis
3
Cycadolepis
2
2
Weltrichia J
-
—
—
—
Leguminanthus
PENTOXYLALES
Lindtheca
3
Taeniopteris
-
-
GNETOPSIDA
NATALIGMALES
Nataligma
3
Gontrigtossa
—
—
ORDERS indet. (2 total)
—
—
Graciliglossa
—
-
-
—
Cetiglossa
-
FRAXINOPSIALES
Fraxinopsis
4
Yabeiella
—
-
-
—
Junqites
—
-
20 genera
27 genera
15 genera
Tab. 13. Affiliated organs in the Molteno, classification
Female/foliage/male affiliates : established in eight cases
Female/foliage affiliates: established in six cases
Foliage/male affiliates: established in two cases
Affiliated organs
20
d^TRELITZI A 15 (2003)
4. MEASURING BIODIVERSITY
Observed diversity (gymnosperms)
'Because they are subject to different controls, the individual
organs ( leaves , pollen, fruit etc.) will tend to supply comple-
mentary information' (Ferguson 1992). In considering the
observed diversity of Molteno gymnosperms, the three organs,
foliage, female fruit and male fruit, are individually and collec-
tively accounted for (Tab. 15). It is assumed, for this purpose,
that the affiliations reflected in Tab. 13 have been correctly
established. Intriguingly, and for unclear taphonomic or mor-
phologic reasons, different organs show greater or lesser levels
of observed diversity within the different orders or classes. A
broad pattern emerges from our work in this regard: the males
appear more diverse in the more primitive groups (Pinopsida),
the females more diverse in certain of the more advanced groups
(e.g. Peltaspermales and Fraxinopsiales), and the foliage more
diverse in the Umkomasiales.
Foliage. 8 classes, 17 orders, 24 families, 27 genera, 113 species
The foliage comprises by a very wide margin the most abun-
dant element of the gymnosperm flora, far exceeding female
and/or male fruit in preserved biomass. With 113 recognised
species, the foliage reflects from two to four times the diversity
seen in either the female or male fruit, but as the higher taxonom-
ic ranks are progressively considered the relative diversity of the
foliage declines.
Female fruit. 8 classes, 18 orders, 18 families, 20 genera, 51 spp.
From generic through to order level, female fruit diversity
in the Molteno outstrips that of the male fruit at a proportion
averaging around 3:2. There is. however, a clear increase in
diversity disparity seen when shifting up through the higher
taxonomic ranks (see text tab., p. 17). From the various classes
included as incertae sedis through to the Gnetopsida (Tab. 15),
the females strongly outnumber the males in observed richness.
Significantly, the males are represented, in this bracket, only in
the Fredlindiales.
Male fruit. 4 classes. 1 1 orders, 13 families, 15 genera, 35 spp.
In the Molteno it is only in the Pinopsida and Cycadopsida that
the males show similar observed diversity to the females, (males,
6 gen., 12 spp.; females, 5 gen., 14 spp.). It is interesting to observe
that while the male pinopsid cones occur exceedingly rarely in the
Molteno (apart from a single prolific palaeodeme of Rissikianthus
from Pen 321), their diversity matches or exceeds that in their
female and foliage counterparts.
Whole-plant. 10 classes, 23 orders, 32 families, 38 gen., 143 spp.
In reflecting whole-plant diversity at species level, the highest
tally amongst the two or three organs for each multi-organ genus
is taken. These tallies are considered to indicate the minimum
numbers of species (MNSs, Tab. 15) in that while the affiliation
between organ-genera might have been satisfactorily established,
the same cannot necessarily be said for all the included organ-
species. In the case of some of the larger genera (e.g. Hamshawvia/
Sphenobaiera/Stachyopitys), in particular, certain ovulate or
microsporangiate species may well occur in the collection without
foliage affiliates.
In most, but not all genera, the foliage reflects the greatest
diversity. While 19 species of Dicroidium are recognised, for
instance, only eight of Umkomasia and three of Pteruchus are
readily distinguished. As outlined above, however, the males in
some instances and the females in others reveal greater diversity
than the foliage.
Building or approaching a real idea of the observed plant
diversity in the Molteno, or any other formation, necessarily
involves the consideration of all plant organs — and necessarily a
systematic focus on establishing the patterns of affiliation between
these organs.
Unaffiliated seeds
This section on the unaffiliated seeds (Tab. 14, pis 149-151) is
included to provide further insight into biodiversity at generic and
order level. Seeds that show no evident affiliation with any of the
described taxa at these higher ranks most probably represent addi-
tional diversity at these levels.
Only a limited proportion of Molteno seeds are considered
here, the criteria for inclusion being:
• no evident affiliation with any ovulate organ has been estab-
lished;
• the seeds show some clearly distinctive features such as shape
or ornamentation (many other more or less nondescript forms
occur in the collection);
• they derive from the 22 TCs that have proved most prolific with
regard to ovulate organs (Tab. 11), i.e. from Aas 411 down
through to Wal 111.
The 10 seed types included are infrequent, mostly occurring in
only one TC, and generally rare, mostly with five or fewer speci-
mens. Some of the forms may possibly occur in other TCs not yet
closely examined for sparse unaffiliated seeds. It is possible that
types 9 and 10 are of animal rather than plant origin. No attempt is
made here to describe and name the material formally.
The ‘species’ of seed recorded are all morphologically very
distinctive, and also clearly different from any of the seeds affili-
ated with the described Molteno fruit. Our conclusion, as noted
above, is that there remain a significant number of higher-order
taxa of female reproductive structures to be discovered. An addi-
tional column for seeds could have been added to Tab. 15 opposite
and the question of affiliations, classification, whole-plant genera
and diversity expanded accordingly.
assemblages
(taphocoenoses)
man-hours
cleaving
genera
Unaffiliated seeds
1 23456789 10
1
Aas 411 Die Sph
512 12
- - 2 40 1 - -31-
2
Umk 111 Die 2spp
400 10
15 - - - 4 - - - 4
3
Lit 111 Dic/Hei
550 7
4 -! -i -; 3 1 30 -i -: -
4
Birlll Sph 2spp
550 6
-| -; -; 6- -[ -' -' -i -\ -
5
Kap 111 Die spp
65 6
-i -f -> -i -' -i -i -■ -1 -
6
Lut 311 Hei elo
50 6
-1 -\ -: -1 -; - -1 -; -
7
Maz 211 Hei/Dic
85 5
8
Mat 111 Dic/dub
65 5
9
Kon 222 Die odo
40 5
10
San 111 Die era
30 5
-i -, -, -i -, -i -i -, -
11
Bir 311 Hei/Sph
2 5
^ ' I i 1 i 1 , i
12
Hla 213 Die elo
60 4
j -i -i n -1 -i -i .
13
Ela 111 Die odo
10 4
14
Cyp 111 Die era
100 3
15
Tel 111 Hei elo
90 3
-1 "i - J 5; - | -| “! " | - J
16
Pen 411 ” ”
70 3
"1 "J. "t "1 -J i * J t
17
” 321 Dic/Ris
35 3
. > . > _ i . ' 1 _ : _ ; _ i .
18
” 311 Hei elo
35 3
_t - 1 -i -l -i -l -i -i -i -
19
Kle 111 Hei/Dic
9 3
- 1 - 1 4 -i - 1 -i -i -i -i -
20
Bir 211 Sph 2spp
7 3
-1 ’I " | "I "I "I "I " 1 “1 -
21
Tin 121
5 3
-| -I -1 -; -! -: -] -| -
22
Wal 111 Die odo
50 2
2; -
Total individuals
19 2 6 51 9 1 30 3 1 4
Total assemblages (TCs)
2: 1 2 3 4 1 1 1 1 1
Tab. 14. Unaffiliated seeds of the Molteno Fm
Taphocoenoses: the 22 TCs listed are those with the greatest
diversity of ovulate fruit (see Tab. 17)
Unaffiliated seeds: included are only the 10 most distinctive
seeds (from 22 TCs) for which we have been unable to
establish any affiliations
Matrix: the number of curated individual seeds
Measuring biodiversity
TRELITZIA 15 (2003)
21
DIVISION
CLASS i ^
ORDER | o
Genus i
Ovulate | u)
“T1
O
5‘
(Q
<D
species
1
in
! .2
1 o
' Q.
Microsporangiate <n
whole-
plant
(MNSs)
PINOPHYTA (gymnosperms) [
1
1
1
1
PINOPSIDA (Coniferopsida) ]
1
DORDRECHTITALES
1
1
Dordrechtites j 3
- | -
— |~z
3
VOLTZIALES
1
i
— ! —
- ! -
Fredianthus \ 1
1
_ i _
— ] _
Lutanthus 1 3
3
Telemachus i 6
Fleidiphyllum > 1
Odyssianthus < 1
6
Clariphyllum \ 1
1 _
1
PINALES (Coniferales) 1
1
1
Rissikistrobus i 3
Rissikia i 2
Rissikianthus , 4
4
— | —
Pagiophyllum j 1
— 1 —
1
ORDERS indet. (2 total) i
1
1
Gypsistrobus ! 1
— | —
1
Avistrobus > 1
— 1 —
1
— ! —
— ! —
Helvetianthus 1
1
CYCADOPSIDA
i
1
1
CYCADALES |
_ 1 _
Pseudoctenis 1 9
Androstrobus ' 2
9
— ! —
Jeanjacquesia , 3
— [ _
3
i _
Ctenis \ 2
— 1 —
2
— i —
Moltenia i 4
— ! —
4
GINKGOOPSIDA
!
i
i
PELTASPERMALES !
i
Peltaspermum j 5
Lepidopterls ' 2
Antevsia ] 1
5
- 1 —
Scytophyllum i 1
— 1 —
1
MATATIELLALES !
!
Matatiella ' 4
Kurtziana ' 1 6
— 1 _
16
- 1 —
De jersey a i 1
Switzlanthus ] 2
2
GINKGOALES
1
1
Avatia i 1
Ginkqoites i 6
Eosteria i 2
6
Paraqinkqo ] 1
— j —
1
HAMSHAWVIALES
1
1
Hamshawvia ' 4
Sphenobaiera \ 9
Stachyopitys ! 6
9
UMKOMASIALES
1
1
Umkomasia i 8
Dicroidium 1 i 19
Pteruchus > 3
19
Fanerotheca J 4
Dicroidium | | —
— | -
4
PETRIELLALES
1
1
Kannaskoppia , 1
Kannaskoppifolia \ 1 0
Kannaskoppianthus \ 4
10
ORDER indet.
'
1
Cetifructus i 1
- 1 —
- ! i
1
CLASSES indet. (5 total)
I
ALEXIALES
, -
l
Alexia j 1
_ 1 _
— 1 —
1
HLATIMBIALES
1
Hlatlmbia \ 1
Batiopteris \ 5
_ ;~zr
5
ORDERS indet. (3 total) '
1
i
i
Hvstricia i 1
— ! —
_ 1 _
Saportaea 1 1
— 1 —
1
— 1 —
Linquifolium i 1
— i —
1
BENNETTITOPSIDA
1
i
FREDLINDIALES
i
Fredllndia j 1
Halleyoctenis ' 3
Cycadolepisl [ 1
3
"
1
Weltrichia 1 i 2
— | —
- | —
Leguminanthus ! 1
1
PENTOXYLALES
1
1
Lindtheca 1
Taeniopteris ! 8
— i —
8
GNETOPSIDA
|
1
1
NATALIGMALES
1
!
Nataliqma 1 1
Gontriqlossa i 1
_ i _
1
ORDER indet.
|
i
— 1 —
Graclliglossa < 1
~ > —
1
- ! —
Cetiqlossa \ 1
— i —
1
FRAXINOPSIALES |
1
Fraxinopsis 1 3
Yabeiella ! 2
_ i _
3
— 1 —
Junqites 1 2
— ! —
2
20 genera i 51
27 genera 1 1 1 3
i
15 genera i 35
143
Tab. 15. Observed diversity of gymnosperms in Molteno Fm.
Female: 8 classes, 18 orders, 18 families, 20 genera, 51 species
Male: 4 classes, 11 orders, 13 families, 15 genera, 35 species
Foliage: 8 classes, 17 orders, 24 families, 27 genera, 113 species
Whole-plant: 10 classes, 23 orders, 32 families, 38 genera, 143 species
MNSs (minimum number of species): diversity recorded for whole-plant
species is a minimum (see text opposite)
Families: excluded from this table in interest of simplification (for complete
classification including families see pp. 54, 55)
Multi-organ genera: 16 recognized in Molteno (see Tab. 12, p.18)
Measuring biodiversity
22
cYtRELITZIA 15 (2003)
The species in palaeobotany
The study of diversity and diversity trends at the level of
species will become more meaningful in proportion to the level of
objectivity attained in palaeobotanical taxonomy. Current levels of
objectivity versus subjectivity can hardly be considered adequate.
This can be understood by examining the state of taxonomy with-
in the genus Dicroidium.
On Dicroidium taxonomy & diversity
As the most prominent (successful) genus in the Gondwana
Triassic, Dicroidium illustrates particularly well the apparently
impenetrable problems involved in resolving species-level taxono-
my in palaeobotany. Two comprehensive taxonomic revisions of
the genus have been attempted over the past two to three decades
and these show little agreement in either the taxa recognised or in
diversity reflected (Retallack 1977; And. & And. 1983). Few more
recent authors have closely followed either taxonomic treatment
especially closely.
Reticulate evolution
In And. & And. (1983, 1989) we concluded that Dicroidium
reflected a highly complex pattern of reticulate evolution rather
than the far clearer patterns of phyletic gradualism or punctuated
equilibria. Our taxonomy of the genus, in accounting for species,
subspecies and formae was an attempt to reflect this complexity.
While we still recognise the highly reticulate pattern, in accor-
dance with our palaeodeme approach, we offer here an optional
taxonomy by raising all subspecies to the rank of species. In doing
so, our Dicroidium taxonomy becomes compatible with the
approach adopted for the remainder of the gymnosperm systemat-
ics in the present volume.
Retallack 1977 3 gen., 23 spp., 25 var. (39 taxa in total)
And. & And. 1983/89 1 gen., 10 spp., 17 subspp., 15 formae (32 taxa total)
This volume 1 gen., 21 spp., 15 formae (32 taxa total)
Taxonomy as in Retallack 1977
genus ] species | variety
1
Dicroidium
i brownii
brownii
2
\
barrealense
3
i dubium
dubium
4
australe
5
tasmaniense
6
] eskense
7
1 gouldii
8
i incisum
—
9
”
j lancifolium
] lancifolium
10
i lineatum
11
| narrabeenense
i narrabeenense
12
' bursellii
13
i nataiense
-
14
! odontopteroides
| odontopteroides
15
i argenteum
16
\ crassum
17
j moitenense
18
i obtusifolium
19
! remotum
20
i pinnis-distantibus
21
; prolungatum
j
22
i radiatum
1 -
23
”
i superbum
!
24
' townrovii
| -
25
i zuberi
i zuberi
26
1
| feistmanteli
27
1 papillatum
28
i
i sahnii
29
Johnstonia
1 coriacea
j coriacea
30
i obesa
31
”
] dutoitii
|
32
i steizneriana
' steizneriana
33
”
! serrata
34
1 trilobita
! -
35
Xylopteris
\ argentina
j
36
] etongata
| elongata
37
i rigida
38
\ spinifolia
! -
39
j tripinnata
i
Tab. 16. Dicroidium taxonomy as in Retallack 1977
Diversity: 3 genera, 23 species, 25 varieties
Taxonomy as in And. & And. 1983, 1989
genus species subspecies forma
Taxonomy as adapted in this volume
1
Dicroidium
coriaceum
! dutoitii ,
D. dutoitii Townrow 1967
2
1 coriaceum ! -
D. coriaceum (Johnston 1887) Townrow 1957
3
crassinervis
crassinervis stelznerianum
D. crassinervis (Geinitz 1876) And. & And. 1983
4
obtusifolium
5
j triiobitum
6
crassinervis
7
sanifotium
8
nondichotoma
-
D. nondichotoma And. &And. 1989
9
odontopteroides ! orbicuioides -
D. orbicuioides (And. & And. 1983) status nov.
10
odontopteroides odontopteroides
D. odontopteroides (Morris 1845) Gothan 1912
11
koningifoiium
”
12
\ lineatum
D. lineatum (Ten. Woods 1883) And. & And. 1970
13
! hlatimbifolium -
D. hlatimbifolium (And. & And. 1983) status nov.
14
dubium
: tasmaniense -
D. tasmaniense (Johnston 1887) And. & And. 1970
15
' switzifolium -
D. switzifolium (And. & And. 1983) status nov.
16
helvetifolium -
D. helvetifolium (And. & And. 1983) status nov.
17
dubium -
D. dubium (Feitsm. 1878) Gothan 1912
18
narrabeenense
D. narrabeenense (Walkom 1925) Jac. & Jac. 1950*
19
zuberi
D. zuberi (Szajnocha 1888) Archang. 1968
20
hughesi*
1
D. hughesi ( Feistm. 1880) Lele 1962*
21
suberbum
superbum townrovii
D. superbum (Shirley 1898) Townrow 1957
22
superbum
23
”
bipinnatum
24
tripinnatum
25
mazenodifolium -
D. mazenodifolium (And. & And. 1983) status nov.
26
elongatum
elongatum remotipinnulium
D. elongatum (Carruth. 1872) Archang. 1968
27
spinifolium
28
rotundipinnuiium
29
elongatum
30
matatifolium 1
D. matatifolium (And. & And. 1983) status nov.
31
”
; argentinum < -
D. argentinum (Kurtz 1921) And. & And. 1970
32
dimorphum j -
D. dimorphum (And. & And. 1983) status nov.
Tab. 17. Dicroidium taxonomy as in And. & And. 1983, 1989 & this volume
Diversity : 10 species, 32 infra-generic taxa total (And. & And. 1983, 1989)
21 species, 32 " ” ” " (this volume)
* species not occurring in Molteno Fm.
Measuring biodiversity
c/tRELITZIA 15 (2003)
23
On Kannaskoppifolia taxonomy & diversity
Kannaskoppifolia is the fifth most prominent gymnosperm
foliage genus in the Gondwana Triassic (see section on
Prominence , pp. 26-29). Of all the Gondwana Triassic genera, it
provides the best case study for exploring approaches to infra-
generic taxonomy and for probing the problem of recognising
observed diversity. The genus, newly described in this volume,
approaches the ideal for a wide range of reasons:
• generic integrity— with its unique foliage (shape and reticulate
venation) with cuticle showing transversely oriented stomata,
there is little doubt as to the generic integrity of the taxon;
• organic attachment— all three organs, foliage, female and male
fruit, are known with certainty through organic attachment;
• morphometries— both foliage and male strobili display obvious
diversity, with convenient, easily measured morphological features;
• frequency and abundance— in that the taxon occurs frequently,
yet not too frequently, and fairly commonly, yet not abundant-
ly, the database remains manageable;
• autecology— the plant occurs in a spectrum of identified habi-
tats, with different ‘species’ characterising each.
• affiliations and conspecificity — the foliage, in 26 of the 100
Molteno TCs, and male strobilus, in 12 Molteno TCs (invari-
ably co-occurring with the foliage), offer the clear opportunity
for analysing conspecificity of organs. (The female strobilus
remains known in only one Molteno TC and is not subject to
similar analysis.)
We discuss aspects of Kannaskoppifolia taxonomy with the
purpose of reflecting on the ‘species’ in palaeobotany. Can we
arrive at a reasonably objective and stable concept of the species?
Is it possible to find an approach that can be followed by all
researchers to arrive at the same result given the same collection of
specimens? How closely will this reflect the ‘species’ in nature?
Kannaskoppifolia species, habit & habitat
We recognise 10 species of Kannaskoppifolia foliage in the
Molteno (And. & And., in prep.). These are recorded in the matrix
table (Tab. 18) below, showing TCs, habitat preferences, frequen-
cy, abundance, dominant associated genera and man-hours cleav-
ing. The female strobilus, from only a single TC, clearly represents
a single species. The male strobilus, with 92 individuals from 12
TCs, is not as diverse morphologically as the foliage and only four
species are recognised (pp. 292, 293).
The frequency of the 10 foliage species is very variable— with
K. sp. E (Pen 311) occurring in 14 of the 26 TCs yielding the
genus; and, at the other end of the scale, K. sp. C (Umk 1 1 1 ), K. sp.
G (Umk 111) and K. sp. I (Kan 112) each occurring in only a sin-
gle TC. Based on a spectrum of clues (And. & And., in prep.), we
consider the Kannaskoppia/ Kannaskoppifolia plant to be a herba-
ceous pioneer— from a slender creeper to a free-standing herb —
appearing in clearings and newly disturbed sites in a range of habi-
tats. It is associated mostly with Dicroidium riparian forest,
Heidiphyllum thicket and fern! Kannaskoppifolia meadows; and
very infrequently with Dicroidium open woodland, Sphenobaiera
lakeside woodland and Equisetum marsh.
As for the species of most extant genera occupying a particu-
lar region or biome, those of Kannaskoppifolia in the Molteno
might each be anticipated to fill a distinctive niche or ecozone.
With extensive and intensive sampling of Kannaskoppia! Kanna-
skoppifolia from the Molteno and with systematic interpretation of
the relevant sites (assemblages, deposition, and taphonomy) it
should be possible to derive a taxonomy that reasonably reflects
reality as it existed in the original Molteno floodplain. We submit
that an objective methodology towards the recognition of infra-
generic taxa— not far removed from practice in extant botany— is
attainable.
species (reference palaeodemes)
CO <o
<N
CO ^
<D (1) £ qj ■*-» CTO qj c
<muoiuu.o 6 i 2
</)(/)(/)
<A
(A (A (A
CA CA (A
23456789 10
Habitat
preference
Frequency
(no. of TCs)
Abundance
(dominant
genera)
2 S
■2 §
n
Umk 111 Die 2spp
Lit 111 Dic/Hei
- 55
1 1
- 1
Dicr. riparian
forest (type 1)
2 of 2 TCs
5 7
1 23
2 1
10 10
Kap 111 Dic/Ris
San 111 Die era
Hla 213 Dicelo
Mat 111 Die dub
2 1 - -
2 4-1
- 17 - 2
Dicr. riparian
forest (type 2)
4 of 8 TCs
50 20 25
90 - _5
89 49 1
89 18 4
10 4
2 1
11 32
20 7
Cyp 111 Die era
Vin 111 Die odo
Nuw 111 Die zub
Boe 112 Die cor
Dicr. open
woodland
4 of 35 TCs
75 - 24
70 4 28
70 30 -
99 - 14
4
Aas411 Dic/Sph
-150
lake margin
1 of 10 TCs
60 30 1 75 24
512
7
1
77
10
20
40
-
-
99
-
2
35
15
9
99
-
2
100
25
.
75
5
11
35
13
-
94
2
3
70
1
-
98
6
7
15
3
-
98
20
2
10
10
-
79
2
3
20
6
-
89
48
58
90
58
29
99
30
3
50
2
-
97
1
25
-
-
75
-
20
2
-
-
10
22
63
30
39 60
-
15
1
10
-
-
-
20
52
14
12
Aas 111 Hei elo
2
13
” 211 ” *
19
14
” 311 ” "
26
15
Pen 311 ” ’’
41
16
" 411 ” ”
80
17
Kan 112 ” "
19
18
Gre 121 ” "
22
19
Win 111 ” "
4
20
Tel 111 ” ”
33
21
Lut 311 ” ”
66
2 5
- 10
1 2
- 2
- 18
- 25
- 34
- 70
- 10
- 22
1 3
2 4
Heidiphyllum
thicket
10 of 24 TCs
30
- 4
- 16
22
Gre 111 Equ sp
Epuis. marsh
1 of 18 TCs
Cal 211 Hei/Ast
Kan 111 Ast spA
Kom 111 Sph/Dic
Kon 211 Ast 2spp
fern IKannask.
meadow
4 of 4 TCs
Total assemblages
Total individuals
4 4 1 5 14 6 1 10 2 1 1
6 19 19 177 3 79 1 12 32 4 1
1 12
50 92
Tab. 18. Kannaskoppia in the Molteno, taxonomy, diversity & habitat
Measuring biodiversity
24
d^TRELITZIA 15 (2003)
Diversity in reference taphocoenosis
Reference
taphocoenosis
Whole-plant genus
(in systematic sequence)
Foliage
gen.! spp
gen
9
spp
gen
Cf
spp
insects
indivs : spp
Aas 311
Hei elo
TelemachusIHeidiphyllum
10 1
11
1
1
2
2
146 !
31
Pen 321
Dic/Ris
RissikistrobusIRissikia
13
18
2
2
3
3
251 !
12
Maz 211
Hei/Dic
PeltaspermumILepidopteris
13 1
19
5
6
3
3
372 r
34
Pen411
Hei elo
MatatiellalKurtziana
10
11
3
3
1
1
9 !
5
Wal 111
Die odo
Avatia/Ginkgoites
8
11
2
2
1
2
4 !
1
Bir 111
Sph 2spp
Hamshawviai Sphenobaiera
22 '
30
7
9
3
5
474 i
99
Umk 111
Die 2spp
UmkomasialDicroidium
37 !
75
10
14
4
5
166 !
42
Kan 111
Ast spA
Kannaskoppial Kannaskoppifolia
5 !
7
2
2
-
-
-
Hla 213
Die elo
HlatimbialBatiopteris
29 ;
43
4
7
2
4
52 r
27
Kon 222
Die odo
FredlindialHalleyoctenis
16
22
5
6
6
9
26 ;
10
Aas 411
Dic/Sph
Lindthecal Taeniopteris
20 '
30
14
16
8
10
129 1
43
Kon 211
Ast 2spp
NataligmalGontriglossa
12 !
11
-
-
-
-
- r
-
Kap 111
Dic/Ris
Fraxinopsisl Yabeiella
15 ;
19
6
6
3
4
178 !
L.
43
Tab. 19a. Reference taphocoenoses selected for 13 whole-plant genera, emphasis on classification
Sequence: TCs arranged according to systematic sequence of genera
Whole-plant genera: all 1 3 established V () pairs included
(it should be noted that 16 whole-plant genera are recognized in total if the cf 0 pairs are added)
Horizontal divisions: group the genera into taxonomic classes
Diversity in reference taphocoenosis
Reference
taphocoenosis
Whole-plant genus
(in diversity sequence)
Foliage
gen. spp
9
gen. 1
spp
gen
cf
spp
insects
indivs| SPP
diversity
grade plants)
Umk 111
Die 2spp
UmkomasialDicroidium
37 !
75
io !
14
4
5
166 ;
42
v. hiqh
Hla 213
Die elo
HlatimbialBatiopteris
29 ;
43
4 I
7
2
4
52 ;
27
high
Bir 111
Sph 2spp
Hamshawviai Sphenobaiera
22 ;
30
7 ;
9
3
5
474 |
99
Aas 411
Dic/Sph
Lindthecal Taeniopteris
20 ;
30
14 1
16
8
10
129 i
43
Kon 222
Die odo
FredlindialHalleyoctenis
i6 :
22
5 r
6
6
9
26 1
10
medium
Kap 111
Dic/Ris
Fraxinopsisl Yabeiella
15 i
19
6 ;
6
3
4
178 ;
43
Maz 211
Hei/Dic
PeltaspermumILepidopteris
13 !
19
5 ;
6
3
3
372 !
34
Pen 321
Dic/Ris
RissikistrobusIRissikia
13 ;
18
2 1
2
3
3
25 i
12
”
Kon 211
Ast 2spp
NataligmalGontriglossa
12 ;
11
- !
-
-
-
- !
-
low
Pen 411
Hei elo
MatatiellalKurtziana
1° ;
11
3 ;
3
1
1
9 !
5
Aas 311
Hei elo
TelemachusIHeidiphyllum
10 1
11
i i
1
2
2
i46 ;
31
”
Wal 111
Die odo
Avatia/Ginkgoites
8
11
2 ;
2
1
2
4 !
1
Kan 111
Ast spA
Kannaskoppial Kannaskoppifolia
5 i
7
2 1
2
- ; -
- ;
v. low
Tab. 19b. Reference taphocoenoses selected for 13 whole-plant genera, emphasis on diversity
Sequence: TCs arranged according to total foliage diversity (genera) in reference TCs
Horizontal divisions: group the genera into diversity grades
Diversity grades
>50 species
v. high diversity
30-49
high
15-29
medium ”
10-14
low
<10
v. low
Tab. 19c. Diversity grades applied for Molteno TCs
Grades: reflect species diversity (of foliage);
based ideally on optimal sampling
Measuring biodiversity
d?TRELITZIA 15 (2003)
25
Preserved diversity
What is the gap between the observed and the preserved diver-
sity in the. Molteno Fm.? What is the gap between the number of
taxa thus far collected and identified in the Molteno and the total
preserved diversity assuming a theoretically comprehensive sam-
ple of every taxon entombed in the formation?
Fitting the generalised inverse Gaussian-Poisson distribution
(GIGP) to the observed frequency distribution of taxa, yields esti-
mates of the corresponding preserved biodiversity (And. & And.
1995; Anderson et al. 1996). Three extrapolations for plants and
insects, following this statistical technique, yielded provocative
figures:
vegetative species (full flora)— 206 observed , 667 preserved ;
ovulate orders (gymnosperms)— 16 observed , 84 preserved ;
insects (full fauna)— 335 observed , 7 740 preserved.
The diversity of vegetative species preserved in the Molteno
appears to be over three times the observed tally. The projected
leaps from observed to preserved ovulate-gymnosperm orders and
of insect species— five times and 20 times respectively— are even
more extreme. Although the Molteno has been unusually well sam-
pled, these data suggest that the uncollected taxa, at all ranks (to
order), far outnumber the collected taxa.
Consider the jump from 206 observed to 667 preserved vege-
tative species in the Molteno as projected in Anderson et al.
(1996). Roughly half of these species are gymnosperm and half
non-gymnosperm. The projection from observed to preserved
species is roughly three- to four-fold. The projection for whole-
plant genera may (?)fairly be assumed to follow a similar propor-
tional increase. The tally rises, then, from 38 whole-plant genera
observed to roughly 125 preserved (and to ca 250 existed).
Though we have sampled 100 TCs, only the tip of the diversi-
ty iceberg appears to have been skimmed. This is a crucial obser-
vation for a variety of reasons. Consider the origin of angiosperms,
for instance. If the earliest stem angiosperms emerged in the Late
Triassic and if they remained a very insignificant element of the
flora, it is more than likely that current levels of sampling would
virtually miss them altogether. The crown angiosperms (non-eudi-
cots including monocots, plus eudicots) in the light of recent mol-
ecular-based cladistics (The Angiosperm Phylogeny Group 1998;
Nandi et al. 1998; Soltis et al. 2000; Savolainen et al. 2000), are
generally accepted as being monophyletic. This reveals nothing,
however, of the stem angiosperms and the nature of their tree
through the Mesozoic. They may well have been quite diverse with
several (perhaps many) major lineages to order level. The initial
stages of their evolution will most likely remain largely unob-
served within the explosive Late Triassic radiation for many years
to come. How many stem-angiosperm lineages (Anderson 1999, p.
53) arose in the Late Triassic that never made it through the Late
Triassic extinction events?
Existed diversity (& the taphonomic factor)
Extrapolations suggest that as many as 667 vegetative species,
84 gymnosperm orders and 7 740 species of insects are preserved
in the Molteno Fm. What diversity of plants and insects actually
colonised the intracontinental Molteno braidplain? How many
plant and insect taxa that existed in the region during Molteno
times were never preserved in the sediments of the formation?
What was the measure of the taphonomic filter? We simply do not
know. What we can be certain of is that only part of the biocoeno-
sis (original community) is ever represented in the thanatocoenosis
(fossil assemblage) (Ferguson 1992).
In And. & And. (1995) we followed a possibly conservative
estimate that around half of all taxa living in the Molteno
Floodplain Biome were never preserved. In round terms the origi-
nal Molteno diversity might read:
vegetative species (full flora)— 2 000 existed
insects (full fauna) — 20 000 existed
In our 1995 paper we compared the projected Molteno diversi-
ty with that recorded in the seven biomes— Desert, Succulent
Karoo, Nama-Karoo, Savanna, Grassland, Forest, Fynbos— recog-
nised in extant southern Africa (Gibbs Russell 1987; Rutherford &
Westfall 1994). The number of species occurring in the core areas
of these biomes varies widely from 497 to 7 316. The Molteno
flora, assuming 2 000 species (existed), was seen to be on a par
with the Succulent Karoo (2 125 species) and the Nama-Karoo
(2 147 species) biomes of today.
Though the Molteno Biome was topographically, edaphically
and climatically simple and therefore included few vegetation
types (ecozones), it apparently supported a diversity at least akin to
similar biomes in the extant world.
Measuring biodiversity
26
d^TRELITZIA 15 (2003)
5. PROMINENCE (colonisation success)
Colonising the Gondwana Triassic (GT)
Wilson (1992, p. 121), in The Diversity of Life, handled the
concept of success thus: ‘ Success in biology is an evolutionary
idea. It is best defined as the longevity of a species with all its
descendants. The longevity of the Hawaiian honeycreepers will
eventually be measured from the time the ancestral finch-like
species split off from other species, through its dispersal to Hawaii,
and finally to that time when the last honeycreeper species ceases
to exist.' This seems altogether too narrow a usage for such an all-
embracing term. The success of an individual or institution or
civilisation is not measured in terms of its longevity alone.
Duration can be only one of several attributes.
The terms prominence and success are applied in our work
synonymously. The prominence of a genus in the Gondwana
Triassic Empire refers to its relative importance or consequence
and is measured as the sum of the five attributes— Frequency,
Ubiquity. Diversity, Abundance and Longevity (FUDAL).
The DUFA concept was introduced in And. & And. (1989) and
is here slightly expanded and shuffled to read FUDAL: an acronym
which is fortuitously appropriate ('feudal' essentially referring to
the holding of land or, alternatively, to the continuous quarrel or
contention between clans).
Vegetative organs
Considering the far wider occurrence of vegetative versus
reproductive parts and the uncertainties concerning the affiliation
of organs, the FUDAL rating system is based exclusively on the
former. While the measure of prominence based on foliage fossils
alone may be imperfect, the formula provides a good approxima-
tion of the success of the original whole-plant genus.
Attributes of success (Gondwana Triassic)
Frequency (F): measure of repetitiveness of occurrence.
The number of subregions (degree squares), of the 85 across
Gondwana yielding Triassic megaplants, from which the genus has
been recorded. The tally is derived directly from the distribution
maps published here, most being reproduced unchanged from our
previous Molteno volumes (And. & And. 1983, 1989).
Ubiquity (U): measure of general range of occurrence.
The number of superregions (continents), of the five making up
Gondwana, from which the genus has been recorded.
Diversity (D): measure of speciation, radiation, variability.
The number of species recognised in the genus for the Gondwana
Triassic (as documented in this volume).
Abundance (A): measure of quantity.
The norm of the abundance figures for the genus in those assem-
blages (only those judged to represent largely the local flora) in
which it occurs. The data are based exclusively on Molteno assem-
blages since clear abundance figures are rarely available for other
formations.
Longevity (L): measure of duration of the lineage.
The duration in number of international standard ammonite bio-
zones between first and last recorded appearances— as plotted on
the stratigraphic figures in this volume. Longevity will probably
prove more effectively measured in millions of years, but this is not
attempted here as our GT stratigraphic base for plotting generic
occurrence still shows only the ammonite biozones (see pp. 7, 45).
Weighting of attributes
As applied here, the five criteria of success are not given equal
weight in the FUDAL formula. This is not through design but is
simply a consequence of there being 85 productive degree squares
across Gondwana, only five extant continents comprising the
supercontinent, and ca 45-50 million years spanning the Triassic.
Should the criteria be given equal weight and, if so, how? In
measuring evolutionary success, should abundance rate higher
than frequency or longevity higher than ubiquity, as turns out to be
partly or wholly the case in our study? The abundance figures,
where only 10 of 27 foliage genera rate 1% or higher, might well
gain greater meaning through adjustment.
No attempt is made in this study to introduce adjustments to
the numbers. The ratings, as recorded, serve sufficiently our pre-
sent purpose.
Unequal or under-sampling (& the taphonomic effect)
Lepidopteris, as currently known, occurs very evenly scattered
across Gondwana, yet is recorded from only 19 of 84 degree
squares, while Dicroidium is recorded from 45 degree squares.
This discrepancy is almost certainly a reflection of under-sampling
or under-reporting. Being far less common than Dicroidium, com-
prising around 1% compared to 90% of assemblages as a norm,
more intensive sampling is needed before finding specimens of the
genus. Lepidopteris is in reality quite probably preserved in as
many degree squares as is Dicroidium. Similar under-scoring
would hold also for other foliage genera.
FUDAL fingerprints
The FUDAL fingerprint (formula) for each genus is clearly
distinctive and, along with ‘geostrat’ occurrence, it tells a great
deal about the kind of parent plant being considered. Dejerseya
(6/2/1/11/2) and Kannaskoppifolia (23/3/10/-/26), for instance,
could hardly be more different in terms of colonisation, diversifi-
cation and autecology. Dejerseya, interpreted as a shrub to small
tree that appeared (apparently) only late in the Triassic, is very
infrequent yet common where it occurs and, though morphologi-
cally variable, never appears to have had the time to diversify.
Kannaskoppifolia, seen as a herbaceous pioneer, appeared early in
the Triassic and colonised widely through Gondwana during the
rest of the period, becoming frequent (though always rare) and
well diversified.
Frequency most closely reflects prominence
Of the five attributes defining prominence, frequency (F) is the
measure that most closely reflects it. In Tab. 20a, where the 26
Molteno gymnosperm genera are ordered according to decreasing
prominence, it is readily seen that they also fall very nearly in cor-
rect sequence following decreasing frequency. The prominence
hierarchy reflects very closely the frequency hierarchy. In view of
this, a first approximation of generic prominence in the Gondwana
Triassic can be derived directly from the frequency values. Or put
another way, frequency might be considered the primary attribute
of prominence or success (at least for the GT) rather than longevity
as proposed by Wilson (1992).
Diversity least closely reflects prominence
In contrast to frequency (F), diversity (D) reflects promi-
nence—at least for Gondwana Triassic gymnosperms — least close-
ly. The diversity value for several genera falls far out of sequence
when the genera are plotted according to prominence. Heidi-
phyllum, with only three species on the one hand, and Kurtziana
and Kannaskoppifolia with as many as 20 and 10 species, respec-
tively, on the other, fall well out of sequence.
Prominence
dA'RELITZIA 15 (2003)
27
Hier-
archy
Molteno
foliage genera
(gymnosperms)
of 84
of 5
spp.
%
m.yrs.
Prom-
inence
grade
F
U
D
A
L
1
Dicroidium
45
5
PI
90%
27
188
5
2
Heidiphyllum
26
5
3
95%
18
147
3
Sohenobaiera
26
3
12
30%
26
99
4
Taeniopteris
32
5
10
2%
20
69
5
Kannaskoppifolia
23
3
10
-
26
62
6
Linguifolium
18
4
5
-
27
56
7
Ginkgoites
21
4
9
-
17
51
8
Lepidopteris
19
4
5
1%
21
50
3
9
Pseudoctenis
16
3
11
3%
14
47
10
Yabeiella
21
3
2
-
17
43
11
Rissikia
17
5
2
1%
14
39
1?
Kurtziana
6
2
20
-
2
30
13
Gontriglossa
10
4
1
1%
9
25
14
Halleyoctenis
7
2
4
2%
9
24
15
Dejerseya
7
2
1
11%
2
23
16
Paraginkgo
5
2
1
-
14
22
2
17
Jungites
3
2
3
-
13
21
18
Batiopteris
7
3
7
-
2
19
19
Saportaea
3
3
2
-
9
17
?n
Moltenia
4
3
4
_
4
15
21
Pagiophyllum
2
2
2
-
6
12
22
Ctenis
4
3
2
-
2
11
23
Jeanjacquesia
3
1
3
-
2
9
24
Clariphyllum
3 1 1 - 1
6
1
25
Graciliglossa
1 1 1 - 1
4
26
Cetiglossa
1 1 1 - 1
4
27
Scytophyilum
1 1 1 1
4
Molteno
foliage genera
(gymnosperms)
of 84
of 5
spp.
%
m.yrs
E§
a> o
i Jo
F
U
D
A
L
e§
o-.E
2
Heidiphyllum
26
5
3
95%
18
147
24
Clariphyllum
3 11 - 1
6
11
Rissikia
17
5
2
1%
14
39
21
Pagiophyllum
2
2
2
-
6
12
9
Pseudoctenis
16
3
11
3%
14
47
23
Jeanjacquesia
3
1
3
-
2
9
22
Ctenis
4
3
2
-
2
11
20
Moltenia
4
3
4
-
4
15
8
Lepidopteris
19
4
5
1%
21
50
27
Scytophyilum
111 - 1
4
12
Kurtziana
6
2
20
-
2
30
13
Dejerseya
7
2
1
11%
2
23
7
Ginkgoites
21
4
9
-
17
51
16
Paraginkgo
5
2
1
-
14
22
3
Sphenobaiera
26
5
12
30%
26
99
1
Dicroidium
45
5
21
90%
27
188
5
Kannaskoppifolia
23
3
10
-
26
62
18
Batiopteris
7
3
7
-
2
19
19
Saportaea
3
3
2
-
9
17
6
Linquifolium
18
4
5
-
27
56
14
Halleyoctenis
7
2
4
2%
9
24
4
Taeniopteris
32
5
10
2%
20
69
13
Gontriglossa
10
4
1
1%
9
25
25
Graciliglossa
111 - 1
4
26
Cetiglossa
111 1
4
10
Yabeiella
21
3
2
<
17
43
17
Jungites
3
2
3
-
13
21
Tab. 20a, b. Prominence, Molteno foliage genera in Gondwana context
Arrangement: genera in order of decreasing prominence Arrangement: genera in classified order
FUDAL fingerprints: emphasizing the diversity of pattern Horizontal divisions: genera grouped into taxonomic classes
Prominence score: ranges vary widely from 188 down to 4
Prominence grade: for discussion of genera in each grade see text
Prominence hierarchy & FUDAL grades
Molteno foliage genera: Only the gymnosperm foliage genera
recorded from the Molteno are accounted for here. Only four addi-
tional genera, mostly localised and rare, are known from elsewhere
in the Gondwana Triassic (And. & And. 1989):
Voltziopsis Pinopsida eastern Australia L. Trias
P achy dermophy llum Ginkgoopsida E. Aus, NZ L-U.Trias
Ptilophyllum Bennettitopsida eastern Australia M. Trias
Zamites
Prominence hierarchy : In Tab. 20a, we list the 27 known Molteno
gymnosperm foliage genera according to prominence hierarchy.
Dicroidium and Heidiphyllum clearly head the list with promi-
nence figures of 188 and 147 respectively, while Graciliglossa,
Cetiglossa and Scytophyilum occupy the tail, with figures of only
4. Dicroidium approaches the theoretical maximum for the
Gondwana Triassic Empire, while the latter genera manifest the
theoretical minimum.
FUDAL grades : As for abundance and diversity (discussed else-
where), we apply five grades in categorising the success or promi-
nence of a genus. These are defined as follows:
Grade
Prominence
(success grade)
FUDAL
score
5
maximum success
>150
4
high success
75-150
3
intermediate success
30-74
2
limited success
15-29
1
minimum success
4-14
On the Molteno genera & their prominence grades
Grade 5: Dicroidium stands head and shoulders above all other
Gondwana Triassic genera in prominence. As currently known, it
is twice as frequent and twice as diverse (aside from Kurtziana) as
its closest competitors. Dicroidium is the overwhelmingly domi-
nant genus in three (riverine forest types 1 and 2, floodplain wood-
land) of the seven primary habitat types recognised in the Molteno
Biome.
Grade 4: Heidiphyllum and Sphenobaiera , the two genera falling
in this category, are the only other gymnospermous taxa dominat-
ing particular habitat types in the Molteno.
Grade 3: The nine Molteno genera (Kurtziana again being an
exception) falling in this category all occur frequently and ubiqui-
tously across Gondwana and spanned nearly half or more of the
Triassic. They ranged in diversity from two to 20 species, but were
mostly rare, only occasionally reaching 1-3% in abundance.
Grade 2: The eight Molteno genera falling here are distinctly less
frequent and ubiquitous than in the previous category. Most are
known from only two regions in Gondwana. Other than
Batiopteris, none apparently diversified with much success. In the
four least prominent of these genera, however, each of the few
available palaeodemes represents a clearly different species.
Abundance is mostly below 1% and longevity spans less than a
third of the Triassic.
Grade 1: The seven genera of this category, from Pagiophyllum to
Scytophyilum, are extremely minor components of the Molteno
flora and beyond. The last three are each known from only a sin-
gle assemblage (the Umkomaas Waterfall locality, Umk III).
Prominence
28
df TRELITZIA 15 (2003)
MOLTENO GYMNOSPERM GENERA
Female
Foliage
Male
9 0 d
SAf
Molteno
9 0 d
Ind
9 0 d
Ant
9 0 d
Aus
9 0 d
PINOPSIDA
Dordrechtites
Telemachus
Rissikistrobus
Gypsistrobus
Avistrobus
✓
✓
Heidiphyllum
Clariphyllum
Rissikia
Pagiophyllum
Fredianthus
Lutanthus
Odyssianthus
Rissikianthus
Hslvetiarithus
Androstrobus
1 2
- 1
- 3
1 1
3 2 4
1 2
2 2
CYCADOPSIDA
Pseudoctenis
Jeanjacquesia
Ctenis
Moltenia
GINKGOOPSIDA
Peltaspermum
Avatia
Hamshawvia
Umkomasia
Fanerotheca
Kannaskoppia
Cetifructus
Lepidopteris
Scytophyllum
Kurtziana
Dejerseya
Ginkgoites
Paraginkgo
Sphenobaiera
Dicroidium
Kannaskoppifolia
Antevsia
Switzianthus
Eosteria
Stachyopitys
Pteruchus
Kannaskoppianthus
- 1
- ✓
- ✓
5 2 1
- 1 -
4 16 -
- 1 2
1 6 2
- 1 -
- 5 ✓
V V V
✓ - -
- ✓ -
4 9
8 19
4 -
1 10
1 -
1 4
- 3 -
✓ ✓ ✓
✓ ✓ -
- 1 1
- 4 -
- 1 -
1 5 ✓
✓ ✓ ✓
✓ - -
- ✓ .
CLASSES INDET.
Alexia
Hlatimbia
Batiopterls
1 -
1 5
ORDERS INDET.
Hystricia
Saportaea
Linguifollum
BENNETTITOPSIDA
Fredlindia
_ Lindtheca_ _
GNETOPSIDA
Nataligma
Halleyoctenls
Taeniopteris
Cycadolepis 1
Weltrichia J
Leguminanthus
1 3 1
- - 2
J_ 8
1 1
- 1
Fraxinopsis
Gontriglossa
Graciliglossa
Cetiglossa
Yabeiella
Jungites
3 2
- 1
- 1
4 3
- 2
2 1
Tab. 21. Molteno gymnosperm genera in Gondwana context, diversity
Molteno genera : arranged to show affiliations
Matrix of table', species diversity as observed in Gondwana Triassic
✓ = presence (diversity unknown)
Prominence
d^TRELITZIA 15 (2003)
29
Colonising the Molteno floodplain
A cursory glance at the table below (Tab. 22) provides a clear
indication that only two genera of gymnosperm, Dicroidium and
Heidiphyllum (apart from the horsetails, particularly the single
genus Equisetum), dominate the Molteno flora. Sphenobaiera is a
clear, yet distant, third in dominance. It is these four genera— three
gymnosperm and one non-gymnosperm — that dominate the major
habitats/ecozones recognised in the Molteno Floodplain Biome.
Prominence hierarchy & FAD grades (Tab. 22)
Prominence: Where generic prominence for the Gondwana
Triassic Empire is scored according to the FUDAL formula, it is
scored for the more restricted Molteno Formation according to the
FAD (Frequency + Abundance + Diversity) formula. The attributes
Ubiquity and Longevity do not apply in the narrower geographic
space and more limited time frame. It has been shown that fre-
quency most closely reflects prominence for the Gondwana
Triassic (Tab. 20a, b) and it is seen to reflect prominence even more
closely for the Molteno (Tab. 22). When the 27 Molteno gymno-
sperm genera are listed in order of decreasing frequency, their
FAD-scores hierarchy parallels the sequence almost exactly.
FAD grades: As for the FUDAL system in the case of the
Gondwana Triassic, we apply a scheme of 5 FAD grades in cate-
gorising the success or prominence of a genus in the Molteno Fm.
Gymnosperm
genera
frequency
abundance
diversity
plant form
preferred habitat
Mo
Prom
FAD
teno
inence
Grade
i
Dicroidium
75
90%
19
shrub to large tree
forest to woodland
184
2
Heidiphyllum
62
95%
1
woody, reed-like
floodplain thicket
158
3
Sphenobaiera
43
30%
9
shrub to med. tree
lake margin
82
4
4
Taeniopteris
38
2%
8
shrub to small tree
forest to woodland
48
5
Lepidopteris
30
1%
2
medium shrub
riverine forest
33
6
Yabeiella
29
750
3
large tree
32
7
Kannaskoppifolia
26
750
10
herbaceous pioneer
wide spectrum
36
8
Pseudoctenis
21
3%
9
cycad-like (small)
forest to woodland
33
9
Rissikia
21
1%
2
large tree
riverine & wetland
24
10
Ginkgoites
19
250
6
shrub to tall tree
floodplain woodland
25
11
Kurtziana
13
150
16
small tree
29
12
Halleyoctenis
10
2%
3
cycad-like
open woodland
15
13
Batiopteris
10
70
5
creeper
wide spectrum
15
14
Linguifolium
9
75
1
herbaceous pioneer
water margin
10
15
Gontriglossa
8
1%
1
10
16
Dejerseya
5
11%
1
shrub or small tree
forest to woodland
17
17
Moltenia
5
55
4
cycad-like (small)
riverine forest
9
2
18
Clariphyllum
3
51
1
shrub
4
19
Jeanjacquesia
3
8
3
cycad-like (small)
6
20
Paraginkgo
2
43
1
shrub
3
21
Ctenis
2
3
2
cycad-like (small)
4
22
Junqites
1
18
2
shrub or tree
3
23
Graciliglossa
1
13
1
slender creeper
2
24
Cetiglossa
1
3
1
herbaceous undergrowth
2
25
Pagiophyllum
1
2
1
tree
open woodland
2
1
26
Saportaea
1
1
1
herbaceous undergrowth
riverine forest
2
27
Scytophyllum
1
1
1
shrub
2
Tab. 22. Molteno foliage genera, frequency, abundance, diversity
Arrangement: genera in order of decreasing frequency, then abundance
Frequency: number of TCs of 100 sampled in Molteno
Abundance: percentage (bold) — estimated norm in TCs yielding the genus (considering
only TCs judged to represent locally growing communities)
individuals (mild) — tally of specimens in curated collection;
rounded off where >50, estimate where >100
FAD score — abundance added only where 1% or more
Grade
Prominence
(success grade)
FAD
score
5
maximum success
>100
4
high success
50-99
3
medium success
20-49
2
low success
3-19
1
minimum success
2
Prominence grades (FAD scores)
Comparison of prominence , Molteno & Gondwana Triassic
The prominence hierarchy of the 27 Molteno gymnosperm
foliage genera as plotted for the Molteno Fm. in particular, or the
Gondwana Triassic in general, is similar, but not identical. The
widest discrepancy in the positions of particular genera between
the two hierarchies occurs where a taxon has a distinct centre(s)
of prominence somewhere in Gondwana. Examples include
Saportaea, ranked 19th in Gondwana and only 26th in the
Molteno, with a clear centre of prominence in Argentina; and
Linguifolium, ranked 6th in Gondwana and only 14th in the
Molteno, with centres of prominence in Chile, New Zealand and
Queensland along the southern coastal margin of Gondwana.
Prominence
30
d/TRELITZIA 15 (2003)
6. THE MOLTENO BIOME
Here we consider briefly the overall floristics and vegetation of
the Molteno Fm„ including both the gymnospermous and non-gymno-
spermous elements. As elsewhere, the principal emphasis is biodi-
versity.
Floristics & diversity (vegetative)
The Molteno flora is the richest known in the Triassic world.
As currently understood, based on a comprehensive taxonomic
study of the collection (part published, part manuscript), the total
vegetative diversity amounts to 57 genera and 206 species. This
richness, at generic and specific level, is made up almost equally
of non-gymnosperms (30 genera, 92 species) and gymnosperms
(27 genera, 114 species).
Minimum: 1 genus, 1 species (as in several assemblages)
Maximum: 37 genera, 75 species (as in Umk 111 Die 2spp)
Average genera per assemblage
(total vegetative) : 6.81 genera
(non-gymnosperm) : 2.49
(gymnosperm) : 4.34 ”
Average species per assemblage (total vegetative)
(non-gymnosperm)
(gymnosperm)
9.2 species
2.65 ”
6.44 ”
Diversity per assemblage (vegetative)
The overall diversity per assemblage— based on vegetative
taxa— varies considerably, ranging from a single genus and species
(in several assemblages) to the uniquely rich Umkomaas site (Umk
111) with 37 genera and 75 species. The average diversity per
assemblage is relatively low, at around seven genera with nine
species. Although the total diversity of gymnosperms and nongymno-
sperms in the formation is roughly equal, there is a marked dis-
crepancy between the two groups when the average diversity per
TC is calculated. At both generic and specific level, the gymno-
sperms (ca 4.3 genera, 6.4 species) outnumber the non-gymno-
sperms (2.5 genera, 2.7 species) around 2:1 in average diversity.
The seven principal habitat types
species per TC
(average)
TCs
counted
Dicroidium riparian forest (mature)
55.5 spp
2 TCs
Dicroidium riparian forest (immature)
22.5 ”
8 ”
Dicroidium open woodland
9.33 ”
15 ”
Sphenobaiera closed woodland
14.1 ”
7 ”
Heidiphyllum thicket
8.0 ”
18 ”
Equisetum marsh
11.0 ”
2 ”
Fern/ Kannaskoppia meadow
7.0 ”
2 ”
Habitat & biodiversity
The wide variation in diversity values is essentially an expres-
sion of the difference in floristic richness between the seven
Molteno habitat types. The mature stage of the Dicroidium riparian
forest is the richest with an average of 55.5 species perTC, and the
Heidiphyllum thicket and Fern IKaimaskoppia meadow the lowest
with eight and seven species respectively. Though the averages
(from data as in Tab. 25) reflect a very uneven number of TCs per
habitat, the figures are considered to be a fair reflection of reality.
Global diversity trends : When the Molteno diversity data are com-
pared with global data (Niklas et al. 1983) through the geological
column, it is obvious there is need for considerable reassessment,
especially where the Triassic is concerned.
Frequency
At generic level, frequency is seen to vary greatly (Tabs 6a, 22,
23), from appearance in only one TC (six gymnosperm genera ) to
as many as 75 of the 100 TCs (Dicroidium). After Dicroidium the
next most frequent genera are Heidiphyllum (62 TCs), Equisetum
(50 TCs) and Sphenobaiera (43 TCs).
Abundance
The five most frequent genera are also the five most abundant
and — apart from the fern IKannaskoppifolia meadows— charac-
terise the primary habitats (ecozones) of the Molteno Biome.
Foliage genera falling in the common to abundant category range
widely in frequency from Taeniopteris in 38 TCs down to
Dejerseya in only five TCs. Around half of all genera fall in the
rare to very rare category, i.e. fewer than 10 and five individuals
respectively in most of the TCs in which they occur (Tabs 6a, 23).
Plant form
The non-gymnospermous components of the flora, as in extant
vegetation, are interpreted as herbaceous plants covering a wide
range of forms. Some Equisetum species, however, reached an
impressive girth and height. The gymnosperms, on the other hand,
also like their extant relatives, are seen as primarily woody. As
interpreted and documented here (Tab. 23), the 27 gymnosperm
foliage genera fall into the following plant-form categories: large
woody shrubs to trees (11 genera), small to medium shrubs (four
genera), cycad-like plants (five genera), herbaceous undergrowth
(five genera), slender creepers (two genera). The proportion of
woody to nonwoody gymnosperms on this count is 20:7.
Preferred habitat
The Molteno, as outlined in more detail elsewhere, was
deposited on an extensive intracontinental fluvial plain. This con-
stituted a relatively simple biome in which seven primary habitats
(ecozones) have been recognised (Caimcross et al. 1995; Anderson
et al. 1998). In spite of this relative uniformity, topographically,
geologically and climatically, the biome supported a remarkably
rich flora (and insect fauna). At generic and specific level we have
explored the question of preferred habitat. From our study of all
taxa across the 100 sampled TCs it is evident for many genera that
each described species occupied its own relatively narrow habitat
range. Excellent examples of this are clear in Kannaskoppifolia.
Batiopteris, Ginkgoites and, on the basis of ovulate fruit,
Matatiella.
Molteno Fm. biodiversity
The sample'. 250 000 specimens from 100 assemblages
Observed (vegetative)
Preserved
Existed
206 species
876 ”
ca 2 000 ”
Extant southern Africa biodiversity
(by far the world’s richest temperate flora)
Observed : 22 211 species
Existing : ca 25 000 ”
nearly 10% of the world total
Extant global biodiversity
Observed : 263 000 species
Existing : ca 300 000
Biodiversity, Molteno versus extant
References : Anderson et al. 1996; And. & And. 1995; Anderson 1999
The Molteno Biome
d^TRELITZIA 15 (2003)
31
CLASS
SUBCLASS
Genera
species
fre-
quency
1
abundance
i
i
i
plant form
1
1
preferred habitat
BRYOPHYTA
i
i
i
i
i
Muscites
1
16
t
i
i
HEPATOPHYTA
i
Marchantites
9
18
to
mosses & liverworts
] damp/shady undergrowth
INCERTAE SEDIS
1
extremely
\
1
Thallites (+2 gen.)
9
6
rare
1
1
LYCOPHYTA
1
l
1
Cylomeia (+1 gen.)
6
7
herbaceous
floodplain wetlands
SPHENOPHYTA (horsetails)
1
1
1
Phyllotheca
5
6
i very rare
1
1
Schizoneura
4
9
; common
horsetails; reed-like,
i riverine and floodplain
2 genera
3
16
| co-dominant
low to high
wetlands
Equisetum
9_
50
monodominant _
1
1
FILICOPHYTA (ferns)
1
1
1
Drepanozamites
1
: 2
very rare
1
riverine forest
11 genera
32
44
i sparse
1
i (varied)
Dictyophyllum
3
46
8
very rare
ferns
! wide spectrum
Asplenites
2
4 _
riverine forest
miscell. (4 gen.)
8_
13
extremely rare _
1
(varied)
PINOPHYTA
l
1
PINOPSIDA (conifers)
1
1
1
Heidiphyllum
2
; 62
monodominant
woody (bamboo-like habit) , floodplain thicket
Clariphyllum
1
1 3
j very rare
: small to medium shrub
| riverine forest
Rissikia
2
21
1 sparse
large tree
! riverine & wetland
Pagiophyllum
1_
1
extremely rare
tree
open woodland
CYCADOPSIDA
l
1
Pseudoctenis
9
21
sparse
1
forest to woodland
Jeanjacquesia
3
3 ~
; cycad-like;
riverine forest
Ctenis
2
18
2
very rare
generally small
Moltenia
4_
! 5 _
1
GINKGOOPSIDA
1
1
1
Lepidopteris
2
30
sparse
medium shrub
riverine forest
Scytophyllum
1
1
extremely rare
small to medium shrub
1
Kurtziana
16
13
rare
small tree
floodplain woodland
Dejerseya
1
! 5
abundant
shrub or small tree
forest to woodland
Ginkgoites
6
65
19
rare
shrub to tall tree
floodplain woodland
Paraginkgo
1
: 2
, very rare
small to medium shrub
riverine forest
Sphenobaiera
9
43
shrub to medium tree
lake margin
Dicroidium
19
75
i monoaommant
shrub to large tree
forest to woodland
Kannaskoppifolia
10_
26
rare
herbaceous pioneer
wide spectrum
INCERTAE SEDIS
1
1
1
Batiopteris
5
10
rare
creeper
wide spectrum
Saportaea
1
6
1
extremely rare
herbaceous undergrowth riverine forest
Linguifolium
1_
9
; sparse
herbaceous pioneer
; water margin
BENNETTITOPSIDA
1
1
1
Halleyoctenis
3
10
cycad-like
open woodland
Taeniopteris
8
38
l: common
shrub to small tree
forest to woodland
GNETOPSIDA
1
1
Gontriglossa
1
8
sparse
herbaceous pioneer
water margin
Graciliglossa
1
1
very rare
slender creeper
riverine forest
Cetiglossa
1
8
; i
extremely rare
i herbaceous undergrowth
Yabeiella
2
29
sparse
large tree
i
Jungites
2_
1 1
very rare
shrub or tree
i
i
Tab. 23. Molteno floristics (vegetative foliage taxa), observed diversity, frequency,
abundance, interpreted plant form & habitat
Non-gymnosperms : 30 genera, 92 species
Gymnosperms: 27 genera, 113 species
Total vegetative: 57 genera, 205 species
Species: based on full taxonomic review of Molteno flora
Frequency: the number of TCs in which the genus occurs
Abundance: based on scale in Tab. 8 & generalised from Tab. 6a
Plant form: as interpreted from all available clues (see And. & And., in prep., sequel to this vol.)
Preferred habitat: as interpreted from all available clues
Unnamed gnetalean (see pp. 14, 18): not included here
The Molteno Biome
32
c/tRELITZIA 15 (2003)
Primary habitats
The seven primary habitats of the extensive Molteno
Floodplain Biome, each supporting a distinctive plant/insect co-
association are typified as follows:
1. Dicroidium riparian forest (type 1): Climax, multistorey, high-
diversity forest lining mature and abandoned channels crossing an
earlier Triassic erosional land surface.
2. Dicroidium riparian forest (type 2): Preclimax, single-storey,
medium-diversity forest lining shifting, braided (occasionally
meandering) channels of the alluvial floodplain.
3. Dicroidium woodland: Low to medium-diversity woodland of
the open floodplain.
4. Sphenobaiera woodland: medium-diversity woodland fringing
lakes in the floodplain.
5. Heidiphyllum thicket: Monodominant to monospecific shrubby
coniferous stands associated with areas of higher water-table in the
floodplain, or on sandbars in the river channels.
6. Equisetum marsh: Monospecific horsetail stands of the flood-
plain marshes and lake margins, or on sandbars in the river chan-
nels.
7. Fern IKannaskoppifolia meadow: Low-diversity herbaceous
colonies occupying the sandbars of the braided river system.
Identifying preferred habitats
A matrix table (Tab. 24) plotting the 27 gymnosperm foliage
genera against the 57 (of 100) TCs yielding fertile genera provides
the basis for interpreting preferred habitats. The taphocoenoses
(TCs) studied in the survey of a fossil flora, however, are not clean
samples of plant communities like the releves in a phytosociologi-
cal survey of an extant flora. They include a variable and incom-
pletely understood mix of taxa from surrounding communities.
With increasing knowledge of taphonomy in general and of the
Molteno flora and ecosystem in particular, clarification of the
autochthonous, parautochthonous and allochthonous elements of
each TC will steadily improve. From the patterns of occurrence
emerging in Tabs 24—28, it is possible to offer preliminary obser-
vations on habitat preferences of the whole-plant genera bearing
the described ovulate structures.
A further consideration is the distinction between species and
genera. It is the species, not genera, that comprise plant communi-
ties—each species within a genus having evolved to fill a different
niche in the environment. Broader comment has been made in the
case of Kannaskoppia (p. 23), a Molteno ovulate structure whose
foliage is known through organic attachment. A systematic study of
the foliage (And. & And., in prep.) reveals several distinct species,
all interpreted as herbaceous pioneers, exploiting different habitats.
Preferred habitats of the Molteno gymnosperm taxa
The 20 Molteno ovulate genera are discussed below in the
sequence followed on Tab. 26, i.e. according to their phytosocio-
logical occurrence in the seven primary habitat types.
1 ) Umkomasia: 22 TCs, ca 500 individuals
Affiliation', with Dicroidium (Grade 4 reliability)
Species diversity: 8
Preferred habitat : Dicroidium riparian forest and woodland. While
Umkomasia occurs frequently and often commonly in the three
Dicroidium- dominated habitats, it is very infrequent and rare else-
where.
Taphonomic filter: A number of woodland and forest TCs, though
dominated by the foliage genus Dicroidium and well sampled (e.g.
Kap 111, Maz 111, Cyp 111), yield Umkomasia extremely rarely or
not at all. Some selective taphonomic filter must be at work. The
fact that Dicroidium foliage often appears commonly in the
Heidiphyllum- thicket TCs, while Umkomasia is virtually absent,
strongly suggests that the two organs, Dicroidium and Umkomasia,
are subject to different taphonomic controls, and are allochthonous
elements in these assemblages.
2) Rissikistrobus: 7 TCs, 85 individuals
Affiliation : with Rissikia (Grade 4 reliability)
Species diversity: 3
Preferred habitat : Dicroidium riparian forest (climax and pioneer)
and less frequently Dicroidium woodland. The link between both
Rissikistrobus and Rissikia (foliage) with Dicroidium- dominated
TCs is very marked; even in the one Sphenobaiera woodland TC
(Aas 411) in which Rissikistrobus occurs, Dicroidium is a domi-
nant element.
3) Hamshawvia : 4 TCs, 24 individuals
Affiliation: with Sphenobaiera (Grade 4/5 reliability)
Species diversity: 4
Preferred habitat: Dicroidium riparian forest (climax), followed
by Sphenobaiera closed woodland.
4) Nataligma: 1 TC, 4 individuals
Affiliation: with Gontriglossa (Grade 3 reliability)
Species diversity: 1
Preferred habitat: Dicroidium riparian forest (climax).
5) Cetifructus: 1 TC, 2 individuals
Affiliation: unknown
Species diversity: 1
Preferred habitat: Dicroidium riparian forest (mature).
6) Alexia: 1 TC, 6 individuals
Affiliation: unknown
Species diversity: 1
Preferred habitat: Dicroidium riparian forest (climax).
7 ) Hlatimbia: 1 TC, 2 individuals
Affiliation: with Batiopteris (Grade 2 reliability)
Species diversity: 1
Preferred habitat: Dicroidium riparian forest (pioneer).
8) Fanerotheca: 27 TCs, ca 250 individuals
Affiliation: possibly with certain species of Dicroidium (Grade 2
reliability)
Species diversity: 4
Preferred habitat: Occurs most abundantly in Sphenobaiera
closed woodland and secondly in Dicroidium riparian forest
(mature and immature). Only single specimens in four TCs have
been collected in Dicroidium open woodland.
9) Fraxinopsis: 18 TCs, 306 individuals
Affiliation: with Yabeiella (Grade 4 reliability)
Species diversity: 3
Preferred habitat: Dicroidium riparian forest (climax and pioneer)
and Sphenobaiera closed woodland. The distribution pattern of
Fraxinopsis is particularly clear in its frequent and common occur-
rence in these habitats and its near absence elsewhere.
Co-occurrence: Fraxinopsis links strongly with Dordrechtites and
Peltaspermum and weakly with Telemachus and Umkomasia. The
linkage with Dordrechtites possibly has to do with both genera
having been wind- or water-dispersed, readily dehisced, conspicu-
ously winged (?)megasporophylls. The occurrence of Yabeiella
(foliage) is markedly wider in the Dicroidium pioneer forest and
open woodland TCs than is Fraxinopsis. Why is the winged seed
often not found in these Dicroidium- dominated TCs, while it is
invariably preserved in the Sphenobaiera- dominated TCs?
The Molteno Biome
C^TRELITZIA 15 (2003)
33
10) Peltaspermum: 17 TCs, ca 250 individuals
Affiliation', with Lepidopteris (Grade 4 reliability)
Specific diversity. 5
Preferred habitat'. Peltaspermum occurs most abundantly in
Sphenobaiera closed woodland TCs, less often in TCs of the vari-
ous Dicroidium habitats, and rarely in those representing
Heidiphyllum thicket. The distribution pattern of the foliage genus
Lepidopteris (found in 30 of the 100 Molteno TCs) points clearly
to the Peltaspermum/Lepidopteris plant being a common (never
dominant), persistent element of the forest and closed woodland
communities of the river bank and lake margin. The discrepancy
between the foliage and fruit distributions can presumably be
traced to their differing taphonomic histories.
Co-occurrence: The pattern of occurrence is most like that of
Fraxinopsis , except in the Dicroidium open woodland TCs where
the two genera are mutually exclusive. Peltaspermum and
Umkomasia , the two most frequently occurring and abundant gink-
goopsid strobili, are often mutually exclusive. The same pattern of
exclusivity is not reflected in the distribution of the foliage affili-
ates— Lepidopteris and Dicroidium — of the two genera. Again,
some taphonomic or seasonal effect must be involved.
11) Dordrechtites: 17 TCs, ca 400 individuals
Affiliation: unknown
Species diversity: 3
Preferred habitat: Dordrechtites occurs more or less equally fre-
quently and commonly in Dicroidium riparian forest (climax and
pioneer), Sphenobaiera woodland and Heidiphyllum thicket TCs,
where the scales are presumed to be allocthonous elements. Their
natural habitat is uncertain. Apart from a few fragmentary strobili,
the numerous specimens occur as dehisced, wind- and/or water-
dispersed, helicopter-blade ovuliferous scales.
Exclusivity: Dordrechtites and Telemachus show a distinctive mutual-
ly exclusive pattern of occurrence. Though each occurs in 17 TCs,
they co-occur (highly disproportionately) in only three TCs.
12) Avatia: 6 TCs, ca 1 10 individuals
Affiliation: Ginkgoites (Grade 2 reliability)
Species diversity: 1
Preferred habitat: Sphenobaiera closed woodland, where Avatia
occurs most abundantly in the Bir 111 and Aas 411 TCs (the best
sampled of the Sphenobaiera woodland TCs).
13) Fredlindia: 3 TCs, 16 individuals
Affiliation: with Halleyoctenis (Grade 3 reliability)
Species diversity: 1
Preferred habitat: Dicroidium woodland and Sphenobaiera wood-
land. The distribution of the likely foliage affiliate, Halleyoctenis ,
corroborates these habitat preferences and points particularly to the
parent plant being a relatively frequent and common component of
the Dicroidium woodland.
14) Lindtheca: 1 TC, 16 individuals
Affiliation: with Taeniopteris homerifolius (Grade 3 reliability)
Species diversity: 1
Preferred habitat: Sphenobaiera closed woodland. This very dis-
tinctive ovulate structure seemingly affiliates with T. homerifolius,
a species that occurs quite frequently (13 TC) and commonly
through the Molteno. The affiliation with Taeniopteris, not conclu-
sive from the Molteno data alone, is based on knowledge of the
order Pentoxylales from the Indian and Australian Lower Jurassic
to Lower Cretaceous. The rarity of Lindtheca compared with
Taeniopteris foliage may be a consequence of the fruit only devel-
oping every several years (as in many cycads today) and then only
for a brief period.
15) Gypsistrobus I TC, 5 individuals
Affiliation: unknown
Species diversity: 1
Preferred habitat: Sphenobaiera woodland.
16) Avistrobus 1 TC, 1 individual
Affilation: unknown
Species diversity: 1
Preferred habitat: Sphenobaiera closed woodland.
17) Hystricia: 1 TC, 1 individual
Affiliation: unknown
Species diversity: I
Preferred habitat: Sphenobaiera closed woodland. The Hystricia
parent-plant, was apparently a very rare and infrequent component
of this lakeside woodland habitat.
18) Telemachus: 18 TCs, ca 300 individuals
Affiliation: with Heidiphyllum (Grade 4 reliability)
Species diversity: 6
Preferred habitat: Heidiphyllum thicket. Telemachus occurs most
frequently and commonly in this category (13 of 18 TCs), far less
frequently in Dicroidium- dominated habitats, and is conspicuous-
ly absent from the Sphenobaiera woodland TCs.
19) Matatiella: 4 TCs, 17 individuals
Affiliation: with Kurtziana, (Grade 2 reliability)
Species diversity: 4
Preferred habitat: Matatiella, though infrequent, appears as a dis-
tinctive species in each of four TCs representing four different
habitats.
Co-occurrence: Matatiella almost invariably links with Kannas-
koppianthus, such that we originally took the two genera to be
female and male of the same plant. The true affiliations of the lat-
ter have since become firmly established.
20) Kannaskoppia
Affiliation: with Kannaskoppifolia foliage (Grade 5 reliability)
Species diversity: 1
Preferred habitat: The pattern of occurrence of the genus Kanna-
skoppifolia and its several distinctive species (And. & And, in
prep.) strongly points to their showing different habitat prefer-
ences. We interpret the species of Kannaskoppifolia as being her-
baceous to twining pioneers occupying sandbanks in the braided-
river, crevasse splays in the proximal floodplain, riparian-forest
clearings or margins, and clearings within the Heidiphyllum thicket
in the floodplain.
The Molteno Biome
34
d?TRELITZIA 15 (2003)
Abundance
NON-
GYMNOSPERMS
GYMNOSPERMS
Pinopsida Cycadopsida
Ginkgoopsida
•2
indet. 'Be. Pe.i Gnetopsida
CJ
assemblages «
(taphocoenoses) §
Dicroidium riparian forest (mature)
Umk 111
Die 2spp
41
26
- -
2
1
7
20 5
- ' 1 -
1 41 ' 1
1 18
-
- -
5 69
42 ' 2
- I _
55' 5 13
3 45 -
Lit 111
Dic/Hei
11.
2
- -
10
10
23
- 40
- i48 6
- i! i
- -
20
4 40
1 50
56 ; -
1 17 1 8
78 1 30 -
- 6 18
Dicroidium riparian forest (immature)
1
i
1
1
1
1
1
Mat 111
Die dub
1
-
1 1
20
7
4
-
- |12 -
- -1 i
- 3
-
25 -
18 89
2 ! -
1
2 ] - -
- 2 -
Hla 213
Die elo
3
2
1 3
11
32
1
- 2
- '27 -
- 7' 1
- 1
-
-
49 89
7 i 2
- 1 -
6 i 8 -
- 1 -
’’ 211
Die 3spp
-
-
-
10
1
-
- 1
- ! 3 -
- 2 1 1
- -
-
- -
7 85
1
-1 2 -
- 1 -
„ 212
-
-
-
1
6
-
- 1
- ! 1 -
- 4 1 1
- -
-
- -
7 91
- 1 -
- 1 1 -
5' 1 -
-
Maz 211
Hei/Dic
2
-
-
7
4
32
1 -
- 1 -
- -i 1
3 -
1 64
- 1 -
- 1 i -
1 1 - -
- 11 -
San 111
Die era
1
-
-
2
1
5
- -
- 1 i -
- -1 1
- 1
-
5 3
- 90
3 ! -
1
2! - -
- 2 -
Kap 111
Dic/Ris
-
-
-
10
4
25
30 38
- 40 -
- 2
- -
-
- -
20 50
6 i -
- 4 i -
3 ' 1 -
- 5 -
Maz 111
Die era
-
-
- -
5
3
5
- 1 -
- -1 2
1 -
12 74
-ii-
1
- 22 -
Dicroidium open
woodland
1
1
1
i
r
i
Kon 222
Die odo
-
-
2 -
1
1
-
- 9
-13 -
- -1 9
-
-
13 -
10 87
1
- - ; 8
4 ! - -
- 1 -
Kra 111
-
-
- 1
-
4
-
- -
- -'10
- 4
-
- -
1 90
- | -
5 1 - -
-
Nuw 111
Die zub
-
-
-
-
4
-
- 2
- ! - -
- -J -
- -
-
- -
30 70
1 1 -
1 1 - -
Boe 112
Die cor
-
-
-
4
-
14
- 12
i
.1 -
- 3
-
-
- 99
1 ' -
_ i _
Mor 111
Die odo
-
1
- 98
2 1 - -
Qua 111
-
-
-
-
-
20
- -
_ 1
- -
-
- -
20 40
1
i
1
- 5 -
Gol 111
Die dub
_ i _
- 99
1 . - -
Cyp 111
Die era
6
1
- -
1
4
24
- 1 -
- 75
83 ]20
2 1 - -
- - -
Ela 111
Die odo
-
-
-
-
4
7
-
- ! 2 -
- -' 5
-
-
3 -
1 87
- 1 -
- i _
1 1 - -
- 6 -
Pen 321
Dic/Ris
-
-
-
32
5
1
- 35
- i 1 -
- -i 9
- -
-
- -
5 51
- ! -
- - 1 5
5 1 - -
- 2 -
Kon 111
Die odo
-
-
- 1
1
12
7
- 2
- 1 5 -
- -| 24
- -
-
- -
5 78
i
- - |10
1 1 - -
- 1 -
Nuw 211
Die 2spp
1
_ 1 _
- 1 -
1 -
3 98
- 1 -
- 1 -
- 1 -
Nav 111
Die odo
1
- 98
- 1 -
Kra 311
5
1
_ 1
- 99
1
- 18 -
Vin 111
Die odo
1
-
-
-
2
28
- -
- ' 3 -
- - ! -
- 7
-
1 -
4 70
2 1 -
- - ! .
4 1 - -
- - -
Boe 111
Dic/Hei
-
-
- -
-
-
42
- -
- 1 7 -
i
■ i
- -
-
- -
- 57
1
i
i
1 1 - -
- 1 -
Sphenobaiera closed woodland
1
1
i
i
1
1
i
i
1
1
Bir 111
Sph 2spp
40
4
7 4
20
90
10
-
- ! i i
- -'70
-
-
43 -
85 1
- ' 1
- - ' 6
70 ' - -
- 40 -
Aas 411
Dic/Sph
-
5
3 50
75
24
1
- 25
2 '19 -
- -1 7
- 15
-
41 -
30 60150 i -
- - 40
1 1 - -
- 75 -
Bir 311
Hei/Sph
-
-
-
-
-
45
- -
_ 1
- -
-
1 -
40 14
1
- - 1 1
1
” 211
Sph 2spp
1
-
-
-
1
3
_ 1 _
- 1 -
3 -
95 4
- 1 -
- 2 ' -
- 1 - -
- - -
Wal 111
Die odo
- 1 2 1
- -1 2
-
-
2 -
3 92
- 1 -
31 - -
- 20 -
Mol 111
Sph pon
1
1
99 12
_ 1 _
_ i _
Tin 121
Sph 2spp
-
-
- -
1
-
4
- 1
- i - -
- -i 1
- -
-
- -
95 1
- 1 -
- - 1 -
2 1 - -
- 1 -
Heidiphyllum thicket
1
1
1
i
1
Gre 121
Hei elo
-
2
-
20
2
98
-
- | -
- | -
-
2
-
- 3
22 1 -
_ , -
Tel 111
-
1
-
48
58
89
- -
1
- -j 1
- -
-
23 -
- 6
33 ' 1
- - 1 -
2' - -
- - -
Kle 111
-
-
-
10
-
90
- -
- 1 - -
- -
-
- -
- -
- ! -
. . ! -
- ! - -
- - -
Lut 4112
Hei/Dic
-
-
-
-
-
50
- -
_ 1
- -
-
- -
- 50
i
i
i
- - -
Aas 111
Hei elo
-
17
-
10
20
77
_ 1 _
- 1 -
1 7
2 1 -
. . i .
2 ' - -
- - -
Kan 112
”
-
-
-
6
7
98
" 1 “
- 1 "
- 1
19 1 -
- 2 1 -
1 1 - -
. _ .
Qac 111
Hei/Dic
1
-
-
-
2
50
1
1
8 50
_ 1 _
_ 1 _
_ 1 -
Aas 611
Hei elo
-
-
-
-
1
80
- 1 -
- 1 -
20
“ 1 -
- 1 "
- 1 "
Lut 111
Hei/Dic
-
-
-
10
-
50
- -
_ 1
- -
-
- -
- 40
1
1
_ 1
- - -
" 511
Hei elo
-
-
-
-
1
80
. 1 .
- 1 -
20
- 1 -
- 1 -
- 1 -
Pen 411
Hei/Dic
1
3
-
2
3
94
-
- 50
-
-
- 13
so ; -
“ 1 -
" 311
Hei elo
6
2
-
5
11
75
i
1
- 25
41 ' -
1
_ 1 .
Kon 111
n
-
-
-
10
-
84
- -
- 1 - -
- - ! -
- 5
-
- -
- 3
- ! -
- - ! -
- ! - -
- - -
Aas 211
”
10
6
-
-
2
99
i
14
19 1 -
i
i
” 311
18
13
-
-
2
99
- 1
_ i _
- 1 -
9 15
26 ' 1
- 1 -
- 1 -
Win 111
1
-
-
2
3
79
- 1 -
- 1 -
10
- 10
4 1 1
■ i -
Lut 311
”
6
1
-
30
3
99
- -
i
- -i 19
- 5
-
- -
29 58
66 ' -
- - 1 -
23 1 - -
- 9 -
Kle 111
Hei/Dic
-
-
-
-
-
49
- 3
- 1 - -
- -i i
- -
-
- -
8 45
- 1 -
- - ! -
5 1 - -
- . -
Ela 112
Dic/Hei
-
-
- -
10
-
30
- -
i
■ i
- -
-
- -
- 60
i
i
i
_ 1
- - -
Equisetum marsh
i
i
i
i
i
i
i
i
1
1
Gre 111
Equ sp
1
-
-
97
1
i
_ i _
1 -
- 2
i ' -
_ i _
. 1 -
- 1 -
Ask 111
-
1 -
50
9
-
- -
. i - .
- - 120
- -
-
- -
1 21
- 1 -
- 1 -
- 1 "
Fern IKannaskoppia meadow
i
1
i
i
1
Kan 111
Ast spA
-
-
-
22 63
10
- 2
5 1 -
Korn 111
Sph/Dic
1
- -
15
1
i
i
_ 1 _
60 39
30 ; -
i _
Uncertain habitat
i
1
1
i
1
Boe 111
Lep sto
-
-
- -
-
2
7
- -
i
- -'90
1
- -
-
- -
- 5
1
-3i-
i
1
- 5 -
Tab. 24. Molteno foliage genera, abundance
Taphocoenoses (TCs): the 57 TCs are arranged as in Tab. 26
Primary habitats: see notes for Tab. 26
Non-gymnosperms: the 6 major groups encountered in the Molteno
Gymnosperm foliage genera: the 27 genera are arranged in classified sequence
Abundance: bold = % estimate made at site
mild = individuals in curated collection (where <1%)
The Molteno Biome
d/TRELITZIA 15 (2003)
35
Species
diversity
assemblages
(taphocoenoses)
NON-
GYMNOSPERMS
GYMNOSPERMS
Muscites
Marchantites
Thallites etc.
lycopods
horsetails
ferns
Heidiphyllum ^
Clariphyllum §
■o
Rissikia S2.
Q.
Pagiophyllum “
Pseudoctenis o
><
Jeanjacquesia a
Ctenis ■§
</>
Moltenia g-
Lepidopteris
Scytophyllum
Kurtziana G>
Dejerseya J-
Ginkgoites 8
Paraginkgo g-
u
Sphenobaiera
Dicroidium
Kannaskoppifolia
Batiopteris
3'
Saportaea g-
Linguifolium
Halleyoctenis jp
Taeniopteris i?
Gontriglossa
Graciliglossa j?
Cetiglossa °
w
Yabeiella a
01
Jungites
Dicroidium riparian forest (mature)
Umk 111 Die 2spp I 1 1 - - 8 19
J _1 1
□till Dic/Hei
2 2
1 1
1 -
1 1
1 -
1___2_
- 1
1 1
1 -
1 2 -
- 1 2
Dicroidium riparian forest (immature)
Mat 111 Die dub
Hla 213 Dicelo
” 211 Die 3spp
" 212 "
Maz 211 Hei/Dic
San 111 Die era
Kap 111 Dic/Ris
Maz 111 Die era
Dicroidium open
Kon 222 Die odo
Kra 111 ” ”
Nuw 111 Die zub
Boe 112 Die cor
Morlll Die odo
Qua 111 " ”
Gol 111 Die dub
Cyp 111 Die era
Ela 111 Die odo
Pen 321 Dic/Ris
Kon 111 Die odo
Nuw 211 Die 2spp
Nav 111 Die odo
Kra 311 ” ”
Vin 111 Die odo
Boe 111 Dic/Hei
1 1
1 1
1 2
1 1
1 -
1 1
1 -
1 - 3
1 - 1
1 - -
1 - -
1 - -
1 - 1
1 - -
1 - -
woodland
- - 1 - 1
- 1 1
- - 1
- 2 2
1 1 1
2 4 -
1 2 -
1 4 1
- 6 1
- 2 -
1 1 -
- 1 -
- 1 3
1 1 -
2 4 -
1 6 -
1 3 -
- 3 -
- 1 -
1 1 1
- 4 -
1 1
- 1
- 1
- 2
- 1
1 1
1 1
Sphenobaiera closed woodland
Bir 111
Aas 411
Bir 311
” 211
Wal 111
Mol 111
Tin 121
Sph 2spp
Dic/Sph
Hei/Sph
Sph 2spp
Die odo
Sph pon
Sph 2spp
Heidiphyllum thicket
1 1
4 -
1 2
1 1
Gre 121
Hei elo
- 1 -
- 1
1
1 ---!--
1
- 1
- - - 1
1 ! -
1
1
Tel 111
- 1 -
- 3
3
1 - - - 1 - -
- - 1 1
1 - - 4
3 ' 1
21 . -
_
Kle 111
- 1
1
Lut 4112
Hei/Dic
1 . . . ' . .
. 1 _
- - - 1
1
1
1
Aas 111
Hei elo
- 2 -
- 1
4
1 - - - ! - -
- - 1 2
1 ! -
- 1 -
1 ! - -
Kan 112
- -
- 2
2
1 - - - ; - -
1
- - - 1
s ! -
- 1 ! -
1 ] - -
- - -
Qac 111
Hei/Dic
1 - -
- -
1
1 - - - 1 - -
- 1 -
- - 1 1
- 1 _
_ 1 .
- 1 -
Aas 611
Hei elo
- .
- -
1
1 - - - | - -
“ 1 “
- 1
- 1 -
Lut 111
Hei/Dic
- 1
i - - . 1 - .
_ 1 _
- - - 1
_ 1 _
1
1
” 511
Hei elo
- -
-
1
1 - - - 1 - -
1 -
- I -
- I -
- 1 -
Pen 411
Hei/Dic
1 1 -
- 3
1
1 - - - j - -
1
1 -
- - - 1
2 ! -
1
1
” 311
Hei elo
1 1 -
- 1
2
1 - - - 1 - -
- 1 -
- - - 2
3 ' -
_ 1 _
- 1 -
Kon 111
- 1
1 - - - ; - -
1 -
- - - 1
- 1 -
“ 1 -
Aas 211
1 1 -
- -
1
1 . . . 1 . .
_ l _
- 1
2 1 -
1
_ 1 _
” 311
1 1 -
-
2
1 - 1 - 1 - -
- I -
- - 1 1
2 1 1
- 1 -
- I -
Win 111
1 - -
- 1
3
1 ---I--
1
- 1
- - - 1
2 ! 1
1
1
Lut 311
1 1 -
- 1
1
1 ......
- - i2
2 -
--23
1 1 -
1 1 - .
- 2 -
Kle 111
Hei/Dic
1 - 1 - ! - -
- - ! 1
- - 1 4
" 1 "
- 1 -
1 ! - -
Ela 112
Dic/Hei
- 1
1 - - - • - -
. 1 .
- - - 1
. 1 -
1
. 1 _
Equisetum marsh
Gre 111 Equ sp I 1
Ask 111 ” ”
- 2 1
2 2 -
Fern IKannaskoppia meadow
Kan 111 Ast spA I - - - - 2
J - 1 J_
Korn 11_1_Sph_/Dic
Uncertain habitat
Boe 1 1 1 Lep sto I
Tab. 25. Molteno foliage genera, species diversity
Taphocoenoses (TCs): as for the set of tables 24-28
Primary habitats', as for the set of tables 24-28
Non-gymnosperms: the 6 major groups as in Tab. 24
Gymnosperm foliage genera: as for Tab. 24 opposite
Matrix: figures = species diversity
contrasting with abundance in Tab. 24
The Molteno Bionie
36
TRELITZIA 15 (2003)
Foliage
Phyto-
sociological
table
assemblages
(taphocoenoses)
man-hours
cleaving
| o genera
Non-gymnos. </>
(D
gymnosperms o
(S'
total spp w
Dicroidium
CL
Sphenobaiera |
Heidiphyllum g
3
Equisetum ST
ferns
Dicroidium riparian forest (mature)
Umk 111 Die 2spp 1 400 1 1 0 1 29 46 75
69
5
7
2
1
Lit 111
Dic/Hei
550
__6_
32
38
50
1
23
10
10
Dicroidium riparia
Mat 111 Die dub
i for
65
5 St
5
(immatur
10 18 28
e)
89
18
4
20
7
Hla 213
Die elo
60
4
15
28
43
89
49
1
11
32
" 211
Die 3spp
4
2
4
13
17
85
7
10
1
” 212
11
2
2
18
20
91
7
-
1
6
Maz 211
Hei/Dic
85
5
4
15
19
64
1
32
7
4
San 111
Die era
30
5
3
16
19
90
-
5
2
1
Kap 111
Dic/Ris
65
6
5
14
19
50
20
25 10
4
Maz 111
Die era
30
2
3
13
16
74 12
5
5
3
Dicroidium open w
Kon 222 Die odo
ood
40
an
5
d
8
14
22
87
10
1
1
Kra 111
20
2
3
6
9
90
1
-
-
4
Nuw 111
Die zub
21
2
1
8
9
70
30
-
4
Boe 112
Die cor
6
2
1
11
12
99
-
14
4
-
Mor 111
Die odo
8
1
1
3
4
98
Qua 111
8
1
-
4
4
40
20 20
Gol 111
Die dub
13
1
-
2
2
99
Cyp 111
Die era
100
3
4
8
12
75
-
24
-
4
Ela 111
Die odo
10
4
1
8
9
87
1
7
-
4
Pen 321
Dic/Ris
35
2
4
14
18
51
5
1
32
5
Kon 111
Die odo
34
2
3
16
19
78
5
7
1
12
Nuw 211
Die 2spp
6
1
-
6
6
98
3
1
Nav 111
Die odo
2
1
-
5
5
98
-
1
Kra 311
13
1
-
3
3
99
-
5
Vin 111
Die odo
10
-
2
8
10
70
4 28
-
2
Boe 111
Dic/Hei
9
1
-
8
8
57
-
42
Sphenobaiera clos
Birlll Sph 2spp
ed
550
/OG
7
dland
14 16
30
1
85 10
20
90
Aas 411
Dic/Sph
512
13
8
22
30
60 30
1
75
24
Bir 311
Hei/Sph
2
5
-
7
7
14 40 45
-
-
" 211
Sph 2spp
7
3
2
8
10
4 95
3
-
1
Wal 111
Die odo
50
2
-
11
11
92
3
Mol 111
Sph pon
4
2
-
4
4
12 99
Tin 121
Sph 2spp
3
3
1
8
9
1
95
4
1
Heidiphyllum thick
Gre 121 Hei elo
et
10
1
3
4
7
3
98
20
2
Tel 111
" ”
90
3
7
13
20
6
-
89
48
58
Kle 111
15
1
1
1
2
-
-
90
10
-
Lut 4112
Hei/Dic
2
1
-
2
2
50
-
50
Aas 111
Hei elo
40
1
7
6
13
7
1
77
10 20
Kan 112
15
1
4
9
13
1
-
98
6
7
Qac 111
Hei/Dic
4
1
2
3
5
50
8
50
-
2
Aas 611
Hei elo
3
1
1
2
3
-
80
-
1
Lut 111
Hei/Dic
2
1
1
2
3
40
-
50
10
” 511
Hei elo
2
1
1
2
3
-
80
-
1
Pen 411
Hei/Dic
70
3
6
5
11
13
94
2
3
” 311
Hei elo
35
3
5
6
11
25
75
5
11
Kon 111
4
2
1
3
4
3
-
84
10
-
Aas 211
35
1
3
4
7
-
-
99
-
2
” 311
100
1
4
7
11
15
9
99
2
Win 111
20
1
5
6
11
10
-
79
2
3
Lut 311
50
6
4
14
18
58
29
99
30
3
Kle 111
Hei/Dic
9
3
-
9
9
45
8
49
Ela 112
Dic/Hei
4
1
1
2
3
60
-
30
10
Equisetum marsh
Gre 111 Equ sp
25
2
5
5
10
2
97
1
Ask 111
14
1
7
5
12
21
1
50
9
Fern IKannaskoppi
Kan 111 Ast spA
a me
30
ad
2
ow
4
3
7
10 22 63
Kom 111
Sph/Dic
10
1
3
4
7
39 60
-
15
1
Uncertain habitat
Boe 111 Lep sto
8
1
2
5
7
5
_
7
_
2
Total TCs
Ovulate genera
S ■-
o SS w
S> 2
•a s
m a: x
15 ® 5 5
2 O I
s
ra a
o .a c
ii it E
£ a a>
.n | c -2 C -S
_ _ . I 1 1 I S I
u.ij.a.Q<*u.-jt3<*:c
x S "P
£ § *
197 4 5 7
51 8 2
16 14 17 10
3 10 11 50
5 63 9 40
5 10 2 15
40 26 1 -
6 27 - -
22 2
14 -
11 -
2 1
- 4
35
1
7 20 14
2 - -
60 47 67 50 66 2 - -
47 44 88 100 38 11 16 5
2 5-61 - - -
111
24 12
12
1 1 - - 1 - - -
- - - 1 -
- - - 5 -
- - - 18 -
---6-
---40-
---15-
7 6 7 50 3
1 1 5 - -
2 - - - -
6 -
2 -
1 -
50 7
17 -
2 -
3 1
22 7 4 1 1 1 1
27 18 17 17 6 3 1 1 1 1
18 4 1
Tab. 26. Molteno ovulate genera, phytosociological table
Taphocoenoses (TCs): 57 of the 100 Molteno TCs have yielded fruit & are listed here
Primary habitats: the TCs are grouped according to the 7 Molteno habitat types; within each habitat
the TCs have been shuffled to highlight phytosociological patterns
Vegetative dominants: bold & mild figures as in Tab. 24
Ovulate genera: the 20 genera are shuffled to highlight phytosociological patterns
figures = individuals in curated collection
The Molteno Biome
d?TRELITZIA 15 (2003)
37
Foliage
Microsporangiate genera
Phyto-
sociological
table
assemblages
(taphocoenoses)
man-hours
cleaving
O genera
non-gymnos. w
T3
gymnosperms g
total spp 5!
1 Dicroidium
dominants
2 E
■° a
5 5 E
■s > a
CO.0'
-c 5 a
a a> er
<0 t HI
ferns
Pteruchus
Antevsia
Rissikianthus
Switzianthus
Helvetianthus
Weltrichia
Leguminanthus
Androstrobus
Stachyopitys
Cycadolepis
Fredianthus
Kannaskopp.
Eosteria
Lutanthus
Odyssianthus
Dicroidium riparia
Umk 111 Die 2spp
n fo
400
esl
10
(mature)
29 46 75
69
5 7
2
1
138
7
8
- ! 19
1
1
1
Lit 111 Dic/Hei
550
.1
6 32
38
50
1 23
10
10
36
4
-
50 6
2
-
-! 4
.
-! 9
.
-
-
Dicroidium ripari;
Mat 111 Die dub
in fo
65
res
5
t (immatii
10 18 28
re)
89
18 4
20
7
84
1 -
-'31
1
1
3
Hla213 Die elo
60
4
15 28
43
89
49 1
11
32
20
_
-i 2
” 211 Die 3spp
4
2
4 13
17
85
7 -
10
1
2
1
i
" 212 ’’ "
11
2
2 18
20
91
7 -
1
6
4
Maz211 Hei/Dic
85
5
4 15
19
64
1 32
7
4
27
18
_
- '26
i
San 111 Die era
30
5
3 16
19
90
- 5
2
1
5
1
'! 5
i
Kap 111 Dic/Ris
65
6
5 14
19
50
20 25 10
4
11
-
25
- 1 35
Maz 111 Die era
30
2
3 13
16
74 12 5
5
3
-
-! 5
i
i
Dicroidium open
Kon 222 Die odo
woo<
40
liar
5
id
8 14
22
87
10 -
1
1
13
2
1
5
- ; 35
10
i
i
_ i
Kra 111 " "
20
2
3 6
9
90
1 -
-
4
30
-1 2
Nuw 111 Die zub
21
2
1 8
9
70
CO
o
-
4
15
1
i
Boe 112 Die cor
6
2
1 11
12
99
- 14
4
-
10
-
1
Morlll Die odo
8
1
1 3
4
98
1
- 1 30
_
Qua 111 ’’
8
1
- 4
4
40
20 20
_ 1 _
i
Gol 111 Die dub
13
1
- 2
2
99
1
Cyp 111 Die era
100
3
4 8
12
75
- 24
-
4
-
-
-
-
-
_
- ! 44
_
i
_
.
_
Ela 111 Die odo
10
4
1 8
9
87
1 7
-
4
3
-i 8
Pen 321 Dic/Ris
35
2
4 14
18
51
5 1
32
5
-
-
30
-
_
_
1 | 2
Kon 111 Die odo
34
2
3 16
19
78
5 7
1
12
1
_ 1 _
i
Nuw 211 Die 2spp
6
1
- 6
6
98
3 1
Nav 111 Die odo
2
1
- 5
5
98
- 1
1
i
.
Kra 311 " ”
13
1
- 3
3
99
- 5
1 ' 8
_ i _
Vin 111 Die odo
10
-
2 8
10
70
4 28
-
2
-
-
-
-
-
.
-! 5
_
_
.
_
Boe 111 Dic/Hei
9
1
- 8
8
57
- 42
_ 1 _
i
Sphenobaiera clo
Birlll Sph 2spp
sed
550
WO
7
adland
14 16
30
1
85 10
20
90
1
-113
1
i
i
5
Aas411 Dic/Sph
512
13
8 22
30
60 30 1
75
24
3
-
15
2 -
-
-
- ' 67
3
2 ! 21
-
1
_
Bir 311 Hei/Sph
2
5
- 7
7
14 40 45
1
1
1
" 211 Sph 2spp
7
3
2 8
10
4 95 3
-
1
Wal 111 Die odo
50
2
- 11
11
92
3 -
- ] 2 1
1
Mol 111 Sph pon
4
2
- 4
4
12 99
Tin 121 Sph 2spp
3
3
1 8
9
1
95 4
1
-! 1
Heidiphyllum thic
Gre 121 Hei elo
ket
10
1
3 4
7
3
- 98
20
2
r
i
i
1
-! 2
Tel 111 ” ’’
90
3
7 13
20
6
- 89
48
58
1
-
-
- -
-
-
-145
-
-1 4
8
-
2
Kle 111 " ”
15
1
1 1
2
-
- 90
10
Lut 4112 Hei/Dic
2
1
- 2
2
50
- 50
_ 1 _
1
Aas 111 Hei elo
40
1
7 6
13
7
1 77
10 20
- ' 2 1
Kan 112 ” ”
15
1
4 9
13
1
- 98
6
7
-
-
-
- -
-
-
-! 5
-
-| 5
-
-
-
Qac 111 Hei/Dic
4
1
2 3
5
50
8 50
-
2
-i 2
Aas 611 Hei elo
3
1
1 2
3
-
- 80
-
1
-
-
-
- -
-
-
-
- ! -
-
-
-
Lut 111 Hei/Dic
2
1
1 2
3
40
- 50
10
_ 1 _
i
" 511 Hei elo
2
1
1 2
3
-
- 80
-
1
-
-
-
- -
-
-
- ! -
-
. I .
-
-
-
Pen 411 Hei/Dic
70
3
6 5
11
13
- 94
2
3
i
-! 4
" 311 Hei elo
35
3
5 6
11
25
- 75
5
11
4
_ i _
Kon 111 ” ”
4
2
1 3
4
3
- 84
10
• i ■
- i -
Aas 211 ” "
35
1
3 4
7
-
- 99
-
2
-
-
-
- -
-
-
- ' -
-
-! i
-
-
-
" 311 ” ”
100
1
4 7
11
15
9 99
-
2
- 1 -
-i 4
12
-
-
Win 111 " "
20
1
5 6
11
10
- 79
2
3
1 -
i
I
Lut 311 ” "
50
6
4 14
18
58
29 99
30
3
-
-
-
-
-
-
-'19
-
- '16
2
2
-
Klein Hei/Dic
9
3
- 9
9
45
8 49
-! 3
- 1 “
Ela 112 Dic/Hei
4
1
1 2
3
60
- 30
10
_ 1 _
1
Equisetum marsh
Gre 111 Equ sp
25
2
5 5
10
2
97
1
1
1
-1 1
1
1
_ 1 .
2
Ask 111 " ”
14
1
7 5
12
21
1 -
50
9
15
- 1 ”
~ 1 -
FernIKannaskopp
Kan 111 Ast spA
ia m
30
eac
2
ow
4 3
7
- 10 22 63
1
1
1
1
Kom 111 Sph/Dic
10
1
3 4
7
39 60 -
15
1
- 1 -
-' 2
Uncertain habitat
Boe 1 1 1 Lep sto
8
1
2 5
7
5
- 7
_
2
_
.
.
_
_
_
1
1
_
1
1
_
_
_
Total TCs
22
5
5
4 1
2
1
2 • 27
3
1 '12
l
4
3
1
Tab. 27. Molteno microsporangiate genera, phytosociological table
Taphocoenoses (TCs): all 57 TCs as in Tab. 26 are included;
only 42 of these yield microsporangiate genera
Primary habitats', as for Tab. 26
Vegetative dominants : as for Tab. 26
Microsporangiate genera : the 15 genera are shuffled to highlight phytosociological patterns
figures = individuals in curated collection
The Molteno Biome
38
d^TRELITZIA 15 (2003)
Species
diversity
assemblages
(taphocoenoses)
Ovulate genera
Pinopsida
Q Q: O
Ginkgoopsida
CO 0) O 3
indet. iBe.Pej Gnet,
-2 | c
^ s ^
■o -q
m c
U.
I &
Microsporangiate genera
Pinopsida Cy.i Ginkgoopsida i Benn
= u, £ * £
£ a C c c
c ■c 'S co
.9 p! *9 5
■2 c (B o r
CO <0 "Tt 2
c s o a a 5
^ M Ul » o. ^
I2T1 £
•S -s e
ro £ -
Dicroidium riparian forest (mature)
Umk 111 Die 2spp
Lit 111 Dic/Hei
11
- 3
- 1
2 - - 2
2 - - 1
1 -
1 1
2 1 -
1 1 1
Dicroidium riparian forest (immature)
Die dub
Die elo
Die 3spp
Mat 111
Hla 213
" 211
" 212 ”
Maz 211 Hei/Dic
San 111 Die era
Kap 111 Dic/Ris
Maz 111 Die era
Dicroidium open
Kon 222 Die odo
Kra 111 " "
Nuw 111 Die zub
Boe 112 Die cor
Morlll Die odo
Qua 111 ” "
Gol 111 Die dub
Cyp 111 Die era
Ela 111 Die odo
Pen 321 Dic/Ris
Kon 111 Die odo
Nuw 211 Die 2spp
Nav 111 Die odo
Kra 311 ” "
Vin 111 ” ”
Boe 111 Dic/Hei
11 - - 2 1
- - - - 3 1
----11
- - - - 1 -
1 - - - 2 1
1 - - - 2 1
1 - - - 1 1
1
2 1 1
1 3 -
- 1 -
- 1
2 1
2 1
1 1
2 -
woodland
- - 1 -
2 1
2 -
- 1
- 1
- 1
1 -
1 -
1 -
1 1
1 1
2 3 -
1 1 -
- 1 -
- 1 -
1 1 -
1 1 1
- 1
1 -
1 1
1 -
- 1
Sphenobaiera closed woodland
Birlll Sph 2spp 1
Aas 411 Dic/Sph 1
Bir 311 Hei/Sph 1
" 211 Sph 2spp
Wal 111 Die odo
Mol 111 Sph pon
Tin J21_ _ §ph 2s_pp
Heidiphyllum thicket
2-1 --2-
2 11111-
- - 1 - 1 1 -
1 - - - - 1 -
1 -
- - - - - 1 -
- - 1 - - 1 -
- - 1
- 1 3 - -
1-311
- - 1 1 -
1
Gre 121 Hei elo
- 1
Tel 111 ” "
- 1
Kle 111 ” ”
- 1
Lut 4112 Hei/Dic
- 1
Aas 111 Hei elo
- 1
Kan 112 ” ”
- 1
Qac 111 Hei/Dic
- 1
Aas 611 Hei elo
- 1
Lut 111 Hei/Dic
- 1
" 511 Hei elo
- 1
Pen 411 Hei/Dic
1 1
” 311 Hei elo
1 1
Kon 111 ” "
1 1
Aas 211 ” ”
1 -
” 311 ” ”
1 -
Win 111 " ”
1 -
Lut 311 ” ”
1 -
Kle 111 Hei/Dic
Ela 112 Dic/Hei
11-1
- - - 1
- - - 1
12-1
- 1 - -
Equisetum marsh
Gre 111 Equ sp I
Ask 111 " "
Fern IKannaskoppia meadow
Kan 111 Ast spA I
Kom 111 Sph/Dic J - - - - -
Uncertain habitat
Boe 111 Lep sto
1 - - - - 1
.... 1 .
Total TCs
1718 7 1 1 117 4 6 4 22 27 1 1 ' 1 1 1 i 3 1 ' 1 18
1 3 1 5 1i 2' 5 4 4 27 2212' 3 2 1
Tab. 28. Molteno reproductive genera, species diversity
Taphocoenoses (TCs): as for Tabs 24-27
Primary habitats: as for Tabs 24-27
Ovulate & microsporangiate genera: arranged in classified sequence
Matrix: figures = species diversity
contrasting with abundance in Tabs 26 & 27
The Molteno Biome
d^TRELITZIA 15 (2003)
39
THE SEVEN PRIMARY HABITATS OF THE MOLTENO BIOME
(Umk 111, Lit 111) (Kap 111)
(Aas 311, Aas 211)
Blattodea
(cockroach)
Odonata
(dragonfly)
Coleoptera
(beetle)
Orthoptera
(cricket)
Homoptera
(bug)
Lepidoptera
(moth, butterfly)
Conchostraca
Proportional abundance (number of individuals),
based on reference TC, e g. Bir 1 1 1
cockroaches : beetles : bugs : dragonflies 3:11:1
(Bir 111, Aas 411)
(Gre 111)
(Kan 111)
The Molteno Biome
MOLTENO GYMNOSPERMS
42
dfr. TRELITZIA 15 (2003)
1. FORMAT OF SYSTEMATICS SECTION
Comprehensive coverage
Reproductive element
In this systematic section we aim to present a comprehensive account
to genus and species level of the gymnospermous strobili of the Molteno
flora.
Vegetative element
In our two previous monographs on the Molteno gymnosperms (And.
& And. 1983, 1989) we focussed on the foliage, touching only marginally
on the fruit. Seven foliage genera (five new) are here added: Scytophyllum,
Kurtziana, Kannaskoppifolia gen. nov., Batiopteris gen. nov., Paraginkgo
gen. nov., Graciliglossa gen. nov. and Cetiglossa gen. nov. With these taxa,
the gymnospermous foliage of the formation, at least to genus, is now also
comprehensively covered.
Quality cut-off. Since our primary emphasis is taxonomic biodiversity, we
have endeavoured to cover all fruiting taxa (specimens) in the collection,
however rare and incompletely preserved. In the Molteno flora there exists
a long tail of scarce elements represented by one or very few specimens
from only one or two TCs. This is the very nature of the challenge. Even
so, we must necessarily define a boundary, though imprecise, as the inclu-
sion/exclusion cut-off. Where a specimen shows at least some unique
interpretable features, allowing a meaningful line drawing or photograph,
it is included and named (see also p. 49).
Seeds'. Only where seeds are considered to affiliate with particular ovulate
taxa are they incorporated into the body of the work. There remain many
other seed types not thus accounted for. The 10 most distinctive of these
appearing in the 22 most fructiferous TCs are recorded, but nowhere
described, in a matrix table (Tab. 14) and plates (pis 179-152). They add to
the sense of remarkable diversity in the Molteno.
Classification'. We follow our own comprehensively revised * Global clas-
sification of the gymnosperms' — included in a sequel to this work
(Anderson et al., in prep.) — throughout the taxonomic account. This is
based, conceptually, on the ovulate organs alone.
Organ versus whole-plant genera : In our Prodromus of South African
megafloras, Devonian to Lower Cretaceous (And. & And. 1985), with
the glossopterids (Ottokariopsida) as a core section, we followed a
policy of recognising whole-plant genera and naming them after the
ovulate fruit where possible. Acknowledging the cautionary arguments
of authors such as Chaloner (1986), we have chosen in this volume to
revert to the traditional approach of describing and naming each organ
independently.
General arrangement
We have kept to a succinct, consistent and comparative layout, with the
most comprehensively known whole-plant genera (e.g. Peltaspermum /
Lepidopteris! Antevsia) as standard. The ovulate genus is covered first, fol-
lowed by the microsporangiate affiliate and lastly the foliage. The photo-
graphic plates close the treatment.
Ovulate & microsporangiate genera
All genera are treated systematically following the same essential plan.
The first column of text adopts a particularly tight, synoptic framework, while
the fields thereafter, from 'Reconstructions' to 'Adaptive radiation', allow for
more varied treatment according to the nature of the material at hand.
Type species
The great majority of the genera and type species described here are
new. Type locality, stratum and author are given for non-Molteno types.
Generic diagnosis ( or generic concept)
We follow a general practice in extant botany where a diagnosis is a
statement of those critical few characters that distinguish a taxon from
adjacent taxa. It may also include geographic (e.g. Gondwana) or strati-
graphic (e.g. Triassic) data to signify intent clearly and to exclude a wide
scatter of uncertain material. The heading ‘Generic concept’ is used for
previously established genera.
Generic characters
The description is intended as a succinct, systematic, comparative,
morphological treatment of the reproductive genera described from the
Molteno. It captures that set of diagnostic features defining the genus and
separating it, in particular, from similar or related genera.
Etymology
The derivation of the name is recorded as a standard procedure.
Global range
First and last appearances are noted following the style used in our
' Global classification (a sequel to this work noted adjacent), which, in
turn, follows 'The Fossil Record 2' (Cleal in Benton 1993).
Gondwana Triassic (GT) occurrence
Considering the remarkable scarcity of gymnospermous fruit in the
Gondwana Triassic literature, this field is more often than not limited to an
entry for South Africa (Molteno). Where the genus is more widespread,
comparative statistics (localities and individuals), for the other continents
(regions, basins) are listed.
Molteno occurrence
A standardised documentation of frequency (F), diversity (D) and
abundance (A) enables ready comparison with other ovulate and microspo-
rangiate genera. Accompanying this is a table scoring absolute and relative
abundance for the top (most productive) Molteno TCs for the particular
genus.
Affiliated organs
The affiliation of organs, given central significance in describing the
Molteno gymnosperms and assessing observed diversity, is everywhere
emphasised. Reliability grades of 1 to 5, as defined on p. 16, are followed.
Classification & comparison
Two subfields are briefly discussed:
Suprageneric classification — Provides justification for placing the genus in
the particular family and order chosen.
Intergeneric comparison (Gondwana Triassic)— Gives an indication of the
distinctiveness of the genus, or of its similarities to the taxonomically near-
est genera within the Molteno and Gondwana Triassic.
Reconstructions
Grade 4 or 5 reconstructions of the full strobilus and its diagnostic
parts— for the type species and in some cases the sister species— are given.
(For an outline of the grading system adopted, see p. 44.) Brief text gives
a sense of the level of confidence in the reconstructions. This differs quite
widely according to the quantity and quality of specimens available.
Intactness of cones
In detailing degrees of cone fragmentation, frequency of in situ seeds
or pollen sacs, and the occurrence of dispersed scales and seeds, a sense of
the proportion of immature, mature and senescent cones preserved in each
taphocoenosis (TC) is given. This in turn provides clues regarding the
taphonomic history from biocoenosis (living community) to taphocoenosis
(death assemblage) and to the autecology of the taxon in question.
Classification ( elaborated )
Where the morphological uniqueness and/or other attributes of a genus
prompt further debate on its systematic position, we have added this field.
Alternative classification schemes are discussed, as are other non-GT fos-
sil or extant genera (in boxes) showing some similarity.
Molteno occurrence ( elaborated )
This field is considered only for those 14 ovulate genera —Telemachus,
Rissikistrobus. Peltaspermum. Matatiella. Avatia. Hamshawvia, Umko-
masia. Fanerotheca. Kannaskoppia. Hlatimbia. Fredlindia, Lindtheca and
Nataligma— where the foliage affiliate has been reasonably established
(Tab. 12). (See further comment under ‘whole-plant genera’ opposite.)
Gondwana Triassic occurrence ( elaborated )
Treatment here is particularly varied depending on (a) the extent to
which the affiliation of organs has been established and (b) the frequency
of occurrence of the genus beyond Africa.
Hypodigm tables (pp. 44, 45) and ‘geostrat’ (geographic/stratigraphic)
maps (p. 45) for the Gondwana Triassic are plotted for those ubiquitous
genera occurring in at least one additional continent beyond Africa.
Evidence for affiliations
As previously stressed, the whole question of affiliations is crucial to
our study of floral biodiversity in the Molteno Fm. A consistent and rigor-
ous system of grading has been employed. The evidence for applying the
grades as given is documented more or less expansively according to the
nature of that evidence.
Format
^TRELITZIA 15 (2003)
43
Adaptive radiation
Observed species diversity, as recognised for the reproductive genera,
ranges from one, for around half of all Molteno genera, to six for
Telemachus and Stachyopitys, and eight for Umkomasia (Tab, 15). Those
features that most evidently vary within the genus and serve to define the
species are emphasised, as are their habitat and stratigraphic level within
the formation.
Species (ovulate & microsporangiate)
As for the genera, we follow a tight comparative format in the systematic
account of the species of gymnospermous fruit.
Holotype
The holotype is employed purely as a nomenclatural formality.
Reference palaeodeme (And. & And. 1985, 1989, pp. 17, 564)
It is the reference palaeodeme in our work that is core to defining the
taxonomic integrity of the species. The most comprehensively sampled
palaeodeme (quantity and quality of individuals) is selected as the refer-
ence palaeodeme for the species.
Sister palaeodemes (And. & And. 1989, pp. 17, 564)
Palaeodemes from the same formation (the Molteno in this case) and
belonging to the same species are referred to as sister palaeodemes. A
selection of the most significant examples— those most clearly adding to
the understanding (morphology, cuticle, affiliations, habit, habitat, divers-
ity etc.) of the taxon — is listed. In view of the rarity of the fruit, the major-
ity of species do not, in fact, enjoy the backing of any sister palaeodemes.
Specific diagnosis— as for genera.
Specific characters
Only those characters found most useful in differentiating between the
species of the genus are recorded. Characters uniting the species within the
genus are not repeated.
Etymology— as for genera.
Comment & comparison
Here the text is confined to comparative discussion, noting diagnostic
characters, habit, habitat, abundance, frequency and other attributes, as rel-
evant, of the Molteno species included within the genus.
Foliage genera
The layout for foliage genera is largely as for the reproductive organs,
except that more emphasis is given to occurrence and less to classification
and species differentiation. The information is drawn largely from And. &
And. (1989).
Gondwana Triassic occurrence
A comparative documentation of the prominence of each foliage genus
is given through their FUDAL (Frequency, Ubiquity, Diversity, Abundance,
Longevity) formulae (pp. 26, 27). This provides a measure of colonisation
success.
Molteno occurrence
Since foliage is by far the most abundantly and frequently preserved of
the macroscopic plant organs, it provides the standard for documenting
Molteno and Gondwana occurrence.
Classification & comparison
This text is generally kept brief, with cross-reference to the ‘ global
classification' in a sequel (Anderson et al., in prep.). There the systematic
relationships between taxa, as currently understood, are seen in the broad-
est context. In some cases, as in Fredlindia/Halleyoctenis , the discussion
here is fuller, going beyond the scope in the later work.
Adaptive radiation
In most instances (exceptions including Kannaskoppifolia and
Batiopteris ), the foliage species were described in And. & And. (1983,
1989). They are not redescribed here.
Whole-plant genera
Ecological treatment of selected whole-plant genera (each on a double-
page spread) is covered systematically in And. & And. (in prep.), a sequel to
this work. Included are those 16 multi-organ genera in the Molteno flora
(Tab. 12) where affiliation between foliage and reproductive genera
(female and/or male) has been established with some degree of reliability.
We find it relevant in view of our focus on floristic reality to provide a
brief preview here of the scope and emphasis in the complementary mono-
graph.
Molteno distribution patterns
A table for each whole-plant genus documents the occurrence of the
affiliated organs (foliage, female and male) for each TC in which they
occur. The TCs are grouped according to habitat type to facilitate compar-
ative analysis of distributions. The patterns of co-occurrence, or the lack
thereof, are seen to vary greatly. In certain instances, the female and male
patterns coincide closely (e.g. Umkomasia/ Pteruchus), in others there is no
correspondence (e.g. Kannaskoppia/Kannaskoppianthus).
Seasonality
We attempt, considering all available clues, to reconstruct the annual
(seasonal) growth cycle of the foliage and fruit (female and/or male) of
each whole-plant genus. The cycle is recorded— in table and pie diagram —
from immaturity through maturity to dehiscence and entombment in the
deposit. A primary aim is to seek explanations for the largely discordant
patterns of occurrence of the various well-established to putatively affiliat-
ed organs.
Reference taphocoenosis
The Molteno taphocoenosis in which the whole-plant genus is best
represented is chosen as reference and is documented following a standard
comparative scheme.
Floral association: The assemblage, documenting both vegetative and
reproductive components, is listed to genus— with absolute or relative
abundance. A summary of its principal characteristics follows.
Faunal association: the assemblage, primarily insects, is likewise docu-
mented—in summary table and text.
Phytosociological interpretation: An assessment of the primary and sec-
ondary plant associations (biocoenoses) as represented in the reference
taphocoenosis, is attempted.
Habit reconstruction: The habit of the plant is depicted in its habitat as
interpreted for the reference taphocoenosis. This is based on clues from
modem analogues, taphonomy and morphology. A similar exercise was fol-
lowed for the voltzialean whole-plant genus Aethophyllum by Rothwell et
al. (2000).
Format
44
d?TRELITZIA 15 (2003)
Pen sketches
We make extensive use of pen sketches to illustrate the Molteno gym-
nosperm flora. They provide the most concise interpretation of the genera
and species described and of the differences and similarities between them.
Reproductive taxa: The general aim, allowing for some flexibility, has
been to illustrate all species from three perspectives: a direct sketch reflect-
ing the holotype; a full reconstruction reflecting the species; and detail
sketches and/or reconstructions reflecting diagnostic parts of the cone/stro-
bilus. In certain instances (e.g. Helvetianthus tintinnabulum, p. 132), addi-
tional specimens from the reference palaeodeme are drawn, while in some
limited cases (e.g. Telemachus , pp. 86, 87) only critical elements of the
holotype have been drawn.
Vegetative taxa: For foliage, the coverage is less comprehensive. Here we
aim generally at a single sketch per species. For diverse genera, such as
Dicroidium or Sphenobaiera, a selection of species provides an impression
only of the total morphological range encountered. For established genera,
the sketches are taken directly from our published monographs (And. &
And. 1983, 1989), but for new taxa or those newly recognised as gym-
nospermous, a new range of sketches has been prepared.
Cuticles : In view of their undoubted diagnostic value at generic level, cuti-
cle drawings have been included where possible.
Magnifications : As for photographic catalogue.
Reconstruction grades (as introduced in And. & And. 1989): All pen
sketches of fossil plants are interpretive to some degree. All reflect the sub-
jective view of the artist and/or author. In order to make the intentions of
the author clear a series of reconstruction grades (R1-R5) was introduced
and defined. The grade of each sketch throughout the volume is indicated.
Rl: no intended reconstruction (catalogue number of individual given).
R2: minor intended reconstruction; correcting and cleaning unnecessary or
ambiguous noise (minor irregularities, distortions, breaks in detail)
due to imperfections of preservation or incomplete preparation; based
on a single specimen (catalogue number given).*
R3: intermediate reconstruction; completing or adding leaflets; based pri-
marily on a single specimen (catalogue number given) but other mem-
bers of the home palaeodeme may be consulted (assemblage code
given).*
R4: extensive reconstruction (composite for palaeodeme); full frond, or
other organ, reconstructed from a composite of specimens from a sin-
gle palaeodeme (assemblage code given).
R5: extensive reconstruction (composite for formation); as in R4 but based
on a composite of specimens from sister palaeodemes (assemblage
codes given).
* Interpretations of fruit outline, diagnostic features, ornamentation etc.
are based on the best preserved or most fully prepared areas of the speci-
men. Observations from other specimens in the palaeodeme may be incor-
porated to support the interpretation. The painstaking reproduction of arte-
fact (fine-scale preservational irregularities or uncleared sediment) is mis-
leading and ambiguous and an abdication of purpose in observation and
extrapolation. A later user of the manuscript (without benefit of the origi-
nal specimens) cannot know which irregularities are more real or unreal.
The holotype or reference specimen is the basis for the sketch.
Photographic catalogue
Scope of catalogue
Included are a total of 152 plates with ca 1 500 photographs. They are
dispersed through the monograph, with the plates illustrating a set of affil-
iated genera (whole-plant) following that group of taxa. The focus is very
largely on the reproductive organs, not the foliage (see further below). New
genera such as Lindtheca or Fredlindia are not favoured with greater cov-
erage than established genera such as Peltaspermum or Umkomasia. In line
with the taxonomic text, a consistent format is followed. As a rule, the more
diverse the genus— the greater the number of Molteno species described—
the greater the number of plates. Thus Stachyopitys (with six species) mer-
its 10 plates, while Lindtheca (with one species) merits only two. Further
factors influencing the level of coverage are morphological complexity,
ornamentation and quality of preservation. The plates and line drawings are
seen as complementary, the former emphasising the reality (authenticity) of
the preserved specimens, the latter our interpretation of those specimens at
specified grades of reconstruction.
Palaeodemes: As in our previous monographs, the plates everywhere
reflect— in arrangement and captions— the palaeodeme approach, rather
than individual specimens, in identifying species.
Reproductive organs: The plates comprise a comprehensive cover of the
ovulate and microsporangiate genera described in this volume.
Foliage: While all leaf genera are given recognition in line drawings with-
in the text, only those few genera (e.g. Kannaskoppifolia , Batiopteris)
described here for the first time are covered, in part, photographically. The
bulk of the leaf genera have been thoroughly illustrated in our earlier
monographs (And. & And. 1983, 1985, 1989).
Dispersed seeds: These are covered either with their parent ovulate organs,
or, for those without established affiliations (Tab. 21), on a separate set of
four plates (pis 149-152) at the close of the systematic section.
Cuticles: Although cuticles have by no means been exhaustively studied
for this volume (pp. 46, 47), they are illustrated where available, distinctive
and instructive.
Magnifications: We adhere to a standard series of magnifications— xl, x2,
x4 (or x5), XlO, X20, X40— as with all line drawings, to facilitate compari-
son between taxa.
Hypodigm
Definition
The hypodigm for a given genus or species includes all the illustrated
individuals (photographs or sketches), appearing in the available literature,
considered here to fall within the taxon in question (And. & And. 1983,
p. 226).
Scope (as applied here for the Gondwana Triassic [GT] gymnospermous
fertile genera)
• A separate hypodigm table is compiled only for those GT fruiting genera
that are found both in the Molteno and beyond.
• The full set of relevant GT literature appearing in the bibliography has
been consulted. This comprises a remarkably limited number of refer-
ences with descriptive taxonomic content. Published and reasonably
accessible unpublished works (e.g. theses, survey reports) are included
(indicated by a plus sign).
• Both photographs and pen sketches of all fruit specimens (hand speci-
mens, not cuticle) are considered. Repeats are included, but are clearly
indicated by an asterisk.
Purpose
The tables have wide applicability and facilitate:
• taxonomic decisions on non-Molteno taxa;
• the plotting of distribution maps for each genus and species;
• the focus of attention on assemblages and palaeodemes;
• assembling statistics on the ubiquity and frequency of taxa.
Notes on format & content of table
The design of the hypodigm tables prepared for each genus remains
essentially similar, with minor changes, to that initiated for Dicroidium in
And. & And. (1983, pp. 74, 81-87) and followed in And. & And. (1989).
The changes are noted and briefly justified below.
Author: The references within each Gondwana continent are included
chronologically, not by region and formation as in our earlier works. This
simplification is because of the far fewer fruit illustrations (as compared to
foliage) and our not attempting to circumscribe palaeodemes beyond the
Format
d^TRELITZIA 15 (2003)
45
Molteno. Illustrations from our own previously published works on the
Molteno are not recorded: these would be superfluous as the best material
is all figured in the present volume.
Subregion (degree square): For a full list of the productive subregions
(degree squares), with their letter and number codes spelled out and their
geographic location in the Gondwana Triassic plotted, see Tab. 2, Map 1
(p. 6).
Formation : The geological formations and number codes (referring to the
Standard Triassic Ammonite Zones) are shown in context in Tabs 3 and 4
(p. 7).
Locality: There still exists in the literature no generally adopted terminol-
ogy or ranking in referring to the spatial occurrence of fossiliferous beds.
In And. & And. (1983, p. 3), we introduced a ‘locality’ classification and
have applied it throughout in our palaeoflora series (see Glossary). The
entries in this column generally provide the most precise ‘locality’ data
available (from all sources), but they will inevitably include ‘localities’ of
various and unknown rank.
Name : The taxonomic names are those applied in the references cited.
Illustrations
• We refer here to plate(s) (pi) and figure(s) (f) as given in the original lit-
erature. (In this we deviate from our earlier usage [And. & And. 1983,
1989] to simplify searching the references.) The number in the left
column refers to the tally of illustrated individuals as best as can be
assessed.
• Where a fertile structure is refigured in a later publication, this is includ-
ed but indicated as a repeat and recorded in the relevant columns by an
asterisk. Here we deviate again from usage in And. & And. (1983,
1989)— where any illustrated individual was included only once— largely
because of the far greater rarity of fruit. A perusal of the hypodigm still
provides an immediate tally of the number of individuals illustrated for
each ‘locality’ and species.
• Hand specimens only are included (cuticle illustrations are not consid-
ered).
Species! intactness matrix
• The species listed are the full set recognised (in this work) for the genus
in the GT. They are grouped as two subsets: those recorded from the
Molteno, and those recorded only beyond the Molteno.
• The illustrated specimens in the GT literature, remarkably few and most-
ly of poor quality, are identified as far as possible with respect to the list-
ed set of taxa (more specifically, the reference palaeodemes defining
them). Identifications are considered either ‘acceptable’ ( 1 ) or 'doubtful'
(?1). (All digits in the matrix refer to numbers of specimens.) The poor-
est (‘very doubtful’), individuals are included under spp. indet. The cri-
teria adopted in defining the limits between ‘acceptable’, ‘doubtful’
(inadequate, insecure) and ‘very doubtful’ remain imprecise. The illus-
trated material currently at hand is, in general, of such a quality that it
cannot bear too close a scrutiny. We await the day, not too distant, when
all the best material in collections around Gondwana will be available in
colour and 3D (virtual reality) to all researchers through digital photo-
graphy on the Internet.
• In view of the tentative nature of many of the specific identifications,
Gondwana distribution maps are plotted only for genera. All specimens
included in the matrix are acknowledged on such maps.
• Intactness. This column, with three subcolumns — ‘intact strobili’, ‘frag-
mentary strobili’ and ‘isolated scales’— is newly added here. The pur-
pose is to provide an easy overview of this revealing aspect of quality.
Intactness is also relevant in taphonomic considerations.
Nomenclature
Taxonomy & nomenclature
In line with our practice applied throughout this volume and in our pre-
vious Molteno monographs, we differentiate clearly between specific taxa
(entities) and names.
The hypodigm & nomenclature
A three-fold procedure is followed. Firstly, those entities within the
Molteno considered here to be morphologically distinct at species level are
recognised in the table. Secondly, any further morphological entities from
elsewhere in the Gondwana Triassic considered sufficiently distinct at
species level are added to the hypodigm. Thirdly, the most useful and
appropriate names, following the IUCN rules of priority, are then tagged to
the specific taxa.
Whole-plant genera
We refer to whole-plant genera by linking the names of the two or
three affiliated organs involved, e.g. Peltaspermum/Lepidopteris/Antevsia
or Fredlindia! Halley octenis. The female-cone name is employed first since
this organ provides the primary basis for classification, the foliage name
second as it is by far the most frequently and abundantly found of the
organs, and the male-cone name last.
‘Geostrat’ distribution maps
The geographic and stratigraphic distribution for the Gondwana
Triassic in general and the Molteno Fm. in particular are plotted on
‘geostrat’ maps for each gymnosperm organ-genus— foliage, female and
male— described from the Molteno. For further detail and explanation on
the maps and stratigraphic table, see pp. 6-19.
Gondwana Triassic (geographic)
The geographic occurrence of genera is plotted at degree-square reso-
lution. This affords a clear view of the currently known ubiquity of the taxa.
It will be readily noted that the spread across the Gondwana supercontinent
is invariably wider for the foliage genus than it is for either the female or
male affiliates. It is the foliage map that gives the sense of the colonisation
success of the parent genus. Distributions are based very largely on published
data (solid colour in ‘locality’ circles), but are occasionally augmented by
other sources of information on available collections or reliable on-site
observations (arrow indicating open ‘locality’ circle, e.g. Dordrechtites , p. 61 ).
Gondwana Triassic (stratigraphic)
For convenience of comparison, we have maintained the same strati-
graphic correlation chart as used in And. & And. (1983, 1989)— although
knowledge of the Triassic globally has advanced significantly over the past
20 years. In particular, the latest UNESCO-IUGS (ICS) 2000 International
Stratigraphic Chart (see below) plots the Triassic as spanning 47 million
years from 250-203 Ma and adopts different standard stages (Induan and
Olenekian) for the Lower Triassic. The correlation of plant-fossiliferous
formations around Gondwana clearly needs updating, but this is not
attempted for our present purpose.
203
220 The Triassic Period
International standards
230
UNESCO-IUGS
240 (ICS) 2000
250
Molteno Fm. (geographic)
Geographic occurrence for each of the organ-genera is plotted for the
Molteno at a narrower ‘superlocality’ (units of 10 km diameter) resolution.
This provides an intermediate-scale sense of frequency of occurrence. Also
plotted on these Molteno maps is a degree-square grid for easy cross-refer-
ence to the Gondwana map.
o
c n
UPPER/LATE
Rhetian
Norian
Carnian
</>
<
MIDDLE
Ladinian
cc
Anisian
H
LOWER/EARLY
Olenekian
Induan
Format
46
d^TRELITZIA 15 (2003)
2. CUTICLES
Significance of cuticles
Cuticular preparations have a significant role to play in palaeobotany
and are applied here in addressing three distinct questions:
Classification-. Cuticle remains strongly under-researched and under-
utilised in grouping fossil gymnosperm genera at order or class level.
Molteno case study: A fine example occurs in the Fredlindiales where the
cuticle, along with other morphology, shows the foliage genus
Halleyoctenis (pp. 344, 345) to clearly exhibit proto-bennettitalean fea-
tures. Several cuticular features, such as the transverse stomata, the elliptic
guard cells and anomocytic (two cells) subsidiary cells, and the oblong
interveinal cells with single papillae, support the comparison of
Halleyoctenis with Laurozamites , an undoubted bennettitalean leaf found
very commonly in the Late Triassic of the USA and Mexico. Other cuticu-
lar and macromorphological features clearly indicate the more advanced
position of Laurozamites in the bennettitopsid clade. This becomes partic-
ularly significant when considering the fairly well-established (Grade 3)
affiliation between Halleyoctenis and Fredlindia , the strikingly singular
female cone from the Molteno— which is thus apparently the earliest
known reproductive organ, with Lindtheca (p. 356) of the Bennettitopsida.
Affiliation : There exists great potential for establishing or confirming the
affiliation between organs through similarities in cuticular features.
Molteno case study: FraxinopsislYabeiella (pp. 372-377) provides a con-
vincing example in that Yabeiella (foliage) and Fraxinopsis (female fruit)
yield cuticle with more or less identical and uniquely characteristic fea-
tures. The nonpapillate amorphous cells with meandering walls together
with the anomocytic (4—6 cells), noncutinised subsidiary cells and narrow-
ly elliptic guard cells to the stomata, characterise both Yabeiella and
Fraxinopsis. Only Jungites (foliage) placed in the same family has similar
cuticle. The affiliation between Yabeiella and Fraxinopsis was previously
well established on the basis of mutual occurrence, but the cuticle adds
powerful confirmation.
Habit & habitat : Robustness of cuticle, differences between upper and
lower cuticle, presence of papillae and/or lappets, and stomatal frequency,
may lend clues to growth form and autecology, including climatic factors
(And. & And. 1983, pp. 52, 53).
Molteno case study: A particularly engaging cuticular feature is found to
characterise the two most diverse and often dominant Gondwana Triassic
foliage genera, Dicroidium (p. 256) and Sphenobaiera (p. 222). Both show
a full range from distinctly linear to broad-leaved species. And in each
there is clearly witnessed a parallel range in the cuticle from fully amphis-
tomatic (narrow-leaved forms), with equal stomatal density in the upper
and lower cuticle, to fully hypostomatic (broad-leaved forms), with high
stomatal density in the lower cuticle but no stomata in the upper cuticle.
The hypothesis, as outlined in And. & And. (1983, 1989) and developed
further in a sequel to the present work, is that the narrow-leaved forms of
both genera occurred as dominant trees in the open woodland of the
Molteno floodplains, while the broad-leaved forms grew as undershrubs in
the closed forest lining the braided rivers.
Molteno sampling
The great majority of the 100 sampled Molteno taphocoenoses (assem-
blages) yield specimens preserved as impressions (occasionally moulds
and/or casts); only a few yield compression material with cuticle. The two
assemblages Little Switzerland (Lit 111) and Umkomaas (Umk 111) have
provided by far the most excellently preserved cuticle and, fortuitously, are
also the most diverse Molteno assemblages, with 39 and 73 vegetative
species respectively.
In Tab. 29, we record those female-fruit genera, with abundance data,
occurring at Lit 111 and Umk 111. Also shown are the supposed foliage and
male-fruit affiliates of these genera, again with abundance data for Lit 111
and Umk 111. The cuticles of all the foliage genera have been fully
described in And. & And. (1989), while those of the female and male fruit
are prepared and described in this volume with a view to answering, where
feasible, the three questions noted above. The table gives a clear indication
of the potential in this regard: fully half ( 10 of 20) of the female-fruit gen-
era are known from at least one of the two localities; a higher tally of
female/foliage affiliates (14 of 20 genera) occurs at Lit 111 and/or Umk
111; while the tally of female/male-fruit affiliates (5 of 20 genera) is
markedly lower.
Where the cuticle of a reproductive genus has been previously studied,
i.e. Rissikistrobus, Peltaspermum, Umkomasia and Pteruchus by Thomas
(1933) and/or Townrow (1960, 1962, 1965, 1967), we have not dedicated
time to repeating the research.
Alternative methodologies
Four methods have been followed here, however cursory, in studying
the epidermal features of the Molteno fruit:
a) Conventional maceration.
b) Acetate peels— compression material, where maceration fails (e.g. for
FraxinopsislYabeiella , pp. 370-383).
c) Acetate peels or direct light microscopic observation— fine-grained
impression material (e.g. Lutanthus ornatus pollen sacs, p. 77, pi. 80).
d) Jed Scanning Microscope— compression or impression material, small
hand specimens, where all else fails (e.g. Dordrechtites. p. 63).
Cuticle features
All descriptions of cuticular morphology follow the terminology out-
lined in detail in And. & And. (1989, pp. 58-61).
Cuticle grade
The quality of the cuticle differs widely for the different genera: from
very poor (no features discernible) in Nataligma to excellent (most features
clearly preserved) in Hamshawvia. A scale of five grades (developed in
And. & And. 1989, pp. 54, 55) is applied.
Preservation grades (as in And. & And. 1989)
1. v. poor— cuticle (&/or mesophyl! tissue) definitely present, but aside
from veins, shows no interpretable structures (cell walls, stomata).
2. poor— minimal interpretable structures (besides veins).
3. fair— intermediate preservation, generally very fragmentary, a few
interpretable features, LM photography passable.
4. good— most features visible, LM and SEM photography suitable.
5. v. good (excellent)— features very clear, large pieces available, LM and
SEM photography admirably suitable.
Notes
(a) The same system of grades is applied to individual specimens and to
assemblages (Tab. 29). In the case of assemblages, the grade given refers
to the best preserved taxa (i.e. ginkgoopsids).
(b) Precision in applying grades is difficult since we are dealing with a
continuous scale involving assorted variables: general preservation, clarity
of specific features, size of cuticle fragments.
Cuticles
^TRELITZIA 15 (2003)
47
Female 9
Lit 111
Umk 111
Foliage
Lit 111
Umk 111
Male C f
Lit 111
Umk 111
1
Dordrechtites
I 50 S2
10 : -
_
1 ,
i
_ i
. i
.
2
-
; - ; -
- ; -
-
- 1
_ )
■ i
- i
-
Fredianthus
- ! - ;
-
! -
3
-
I • 1 -
- i .
-
; - 1
_ i
- i
-
Lutanthus
- 1 - ]
-
1 -
4
Telemachus
! - ■ -
- | -
Heidiphyllum
23%
S3:
7%
S3
Odyssianthus
- i - ;
-
I -
5
-
! - ! -
- i -
Clariphyllum
| - i
1
' !
20
SI
-
- 1 - ;
-
I -
6
Rissikistrobus
; io -
45 i *
Rissikia
40
SI
5%
S3
Rissikianthus
- 1 - ;
8
i -
7
-
I - ! -
. ! .
Pagiophyllum
- 1
1
~ 1
- !
-
-
- 1 - ;
-
|
8
Gypsistrobus
; - ! -
- ! -
-
; - i
1
" 1
- !
-
-
- 1 - ;
-
-
9
Avlstrobus
- ! -
-
1 - i
1
“ 1
- i
-
-
- 1 - 1
-
-
10
-
- ! -
-
_[ ' i
1
. '
-
Helvetianthus
6/5
-
1 -
11
-
Pseudoctenis
48
/ 2
i%
SI
Androstrobus
-1
-
-
12
-
- ! -
Jeanjacquesia
6
S3
- !
-
-
- ! - ;
-
-
13
-
; - ! -
- ! -
Ctenis
. . ;
_ 1
1 '
S 2
-
- 1 - ]
-
i -
14
-
- ! -
Moltenia
1 i
SI |
41 !
S3
-
. 1 - ;
-
1 -
15
Peltaspermum
12
17 *
Lepidopteris
1%
s 5 ;
i%
S 4
Antevsia
4
7
i -
16
-
1 - ! -
- 1 -
Scytophyllum
, 1
_ 1
1 '
S3
-
- ! - ;
-
i -
17
Matatlella
- ! -
Kurtziana
1
“ 1
18
S3
-
- i -
-
i -
18
-
Dejerseya
20%
/ 5 |
- !
-
Switzianthus
50 /5
-
! -
19
Avatia
- 1 -
Ginkgoites
44 !
S5 |
- ;
-
Eosteria
- 1 - 1
-
i -
20
-
- ! -
Paraginkgo
40
S5 i
-
-
- 1 - 1
-
i -
21
Hamshawvla
2 / 4
7 S3
Sphenobaiera
1%
/ 5
5%
S 5
Stachyopilys
4 - i
19
! -
22
Umkomasia
50
;ioo *
Dicroidium~~\
50%
S 5 69%
S 5
Pteruchus
35 - ;ioo
: *
23
Fanerotheca
; 3 | -
30 i/3
Dlcroidium J
1 _ 1
1
- i
-
-
- 1 - ;
-
I -
24
Kannaskoppia
, 1
Kannaskoppifolia 51
/ 4
42
S 4
Kannaskoppianthus
9 S3 :
-
! -
25
Cetifructus
2 -
-
1 _ 1
_ 1
- i
-
-
■_[ .-j
-
1 -
26
Alexia
1
6 S3
-
1 '
1 “ 1
1
- i
-
-
-
i -
27
Hlatimbia
, 1
Batiopteris
1 1
1 ' 1
_ 1
2 ;
S2
-
- !
-
i -
28
Hystricia
, 1
-
1 _ I
1
- :
-
-
- 1 - ;
-
! -
29
-
, I
Saportaea
i ;
S2 ;
- ;
-
-
■ i ' ■
-
I -
30
-
, 1
Linguifolium
1 17 :
S3
- i
-
-
- 1 - i
-
i -
31
Fredlindia
, 1
1
Halleyoctenis
8
S3 ;
i :
SI
Cycadolepis
- i
-
; -
32
, 1
i 1
1 " 1
1
-
Weltrichia
1 ' - i
-
' -
33
-
, _ 1
1
-
1 1
1 " 1
I
-
Leguminanthus
- 1 - ;
-
> -
34
Lindtheca
, _ 1
I
Taeniopteris
78
S3 \
_55J
SI
-
-
i -
35
Nataligma
, 1
, " 1
3 i -
Gontriglossa
30 :
S3 ;
5%
S2
-
- ; -i
-
; -
36
-
, _ 1
1
Graciliglossa
1 _ 1
1 1
13 1
/I
-
'1 ' '
-
; -
37
-
| _ 1
1
Cetiglossa
1 _ 1
1 ~ 1
1
3
S 1
-
- ; - i
-
] -
38
Fraxinopsis
10 S3
12 i -
Yabeiella
6
SI
45
S3
-
- 1 - 1
-
| -
39
-
, 1
, “ 1
1
Jungites
18
S3
- ;
-
-
- ! - i
-
1 -
20 genera
! 7 1
10
1
27 genera
; 19 1
1
1
19 ;
15 genera
L
7 ;
4
Tab. 29. Cuticle potential in the Molteno (Lit 111 & Umk 111)
Genera: in classified order reflecting affiliations (see Tab. 13, p. 19)
Female: 10 of 20 Molteno ovulate genera occur at Lit 111 &/ or Umk 111
Male: 8 of 1 5 Molteno male
Foliage: 26 of 27 Molteno foliage
In the locality columns, the first figure gives the number of curated specimens
(or % abundance in the case of foliage) whilst the second gives cuticle grade.
Bold figures: % estimate made at site
Mild figure: individuals in curated collection (where <1%)
/: cuticle (grade 1 — 5 preservation) prepared by HMA (this vol., or our previous vols.)
* : cuticle prepared by Townrow (1960, 1962, 1965, 1967) &/or Thomas (1933)
Cuticles
48
d/ TRELITZIA 15 (2003)
3. TAXONOMIC GUIDELINES
Like justice, palaeobotany is an inexact science. And, as injustice, it is
necessary to seek terms and concepts to reflect the inexactness and to guide
our methodology towards reaching the best approximation of the truth.
Below follows a scheme of the guidelines adopted— much of which might
sound self-evident, yet is not so. Divergent views on palaeobotanical—
indeed all palaeontological— taxonomy, at all ranks from class to species
(and below), have coloured our science for decades and remain extreme.
There is no agreed methodology, there are few agreed guidelines. Nor is
there any sense that current differences might be narrowing. If we are to
track biodiversity patterns through geological time successfully, conver-
gence of approach will be a key necessity.
Seeking the holistic truth (reality): the primary guiding principle
• 'Deep ecological awareness recognizes the fundamental interdepen-
dence of all phenomena . . .’— Fritjof Capra, The web of life (1996).
• What was the holistic reality of life— plants, animals, landscape, cli-
mate-back in Molteno times in the Late Triassic (ca 200 Ma)?
• Taxonomic entities, at all levels, from species and genera to families,
orders and classes, are defined with the biological reality always upper-
most in mind.
• Form taxa, basket taxa, morphospecies and related pragmatic concepts
are therefore avoided wherever possible.
Biodiversity, paleoecology, biogeography, phylogeny
• These are the themes that drive our Molteno research.
• Taxonomy provides the fundamental data essential to exploring these
themes.
• Consistent and rigorously derived taxonomy will underscore sound
hypotheses concerning biodiversity and paleoecology.
• Inconsistent, variously derived taxonomy, will undermine hypotheses
concerning biodiversity and paleoecology.
Nothing by mere authority (‘ Nullius in verba’)
• This is the motto of the British Royal Society, instituted in England in
1660 to promote scientific research and discussion.
• The motto encourages observation and experiment before precedent and
authority.
• Newer, more complete data supersede historic, less complete data.
• Resolution of a dispute is often best resolved by collecting more data.
Beyond our time & space
• To maintain objectivity, to build in hindsight, visualise making taxo-
nomic and related decisions from a vantage point outside your current
framework of reference in time and space. Imagine yourself looking
back critically at your own work— not as a peer reviewer, but as a
descendent-generation reviewer— from 100 years in the future and from
the opposite hemisphere.
• Consider those making taxonomic decisions early in the 20th century
doing likewise. How might this have coloured their approach and their
results?
Further on holism & the interdependence of all phenomena
• The ‘ Unity of knowledge' (Wilson 1998) is the framework in which the
Molteno details are assessed.
• All clues are relevant and synergistic.
• ‘Holistic thinking’ is not ‘circular thinking’.
Contextual thinking
• When studying the taxonomy of the Molteno, we are fully aware that
temporally we are exploring life within the Late Triassic and not, for
instance, the Late Devonian or Late Permian. And we are cognisant that
geographically we are dealing with Gondwana and not Pangaea as a
whole.
• We have in mind from our own previous work (And. et al. 1996, 1999a)
that we are very probably working within a time of peak diversity; near
the height of the ‘Triassic explosion of diversity’, and very close to, if
not coinciding with, the ‘heyday of the gymnosperms’.
Superficial similarities at generic & higher level
• Is a Glossopteris- like leaf found in the Late Triassic a Glossopterisl Is a
Ginkgo-like leaf from the same strata a Ginkgo, or a Podozamites-\ike
leaf a Podoz.amitesl
• This is a perennial problem and a crucial one— involving both morpho-
logical (including affiliations) and contextual considerations.
• Consider Gontriglossa (p. 364) and Glossopteris , for example: contex-
tually, such taxa are less likely to be congeneric, confamilial or conordi-
nal if separated in time by one or more of the five major global extinc-
tion events, the end-Permian in this case.
Superficial similarities at specific & generic level
• Are three widely distinct Glossopteris- like species in the Late Triassic
congeneric or more likely representatives of three distinct genera (see
Gontriglossa, Graciliglossa and Cetiglossw, pp. 364—369)?
• This, again, is a constant problem and one that demands a solution
involving both morphology and context.
• Were these taxa from the Gondwana Permian, they would automatically
be considered congeneric, i.e. Glossopteris.
• Contextually, such taxa are less likely to be congeneric if found in the
Late Triassic (at the peak of diversity) than if found in the Early Triassic
(at the trough of diversity).
• Proof of congeneric status will most likely depend on well-established
fruit (female and/or male) affiliations, and/or clear and unique cuticular
conformity; and/or obvious similarity in mode of attachment.
Habitat & conspecificity
• The recognition of habitats/ecozones within the Molteno introduces a
further parameter in species-level taxonomy.
• A taxonomic law that generally holds in the extant world is that distinct
species within a genus occupy distinct habitats; or conversely, any one
habitat is unlikely to harbour more than one species of the same genus.
• Palaeodemes of a particular genus deriving from a number of clean/un-
contaminated/unambiguous TCs representing a distinct habitat are more
likely than not to represent a particular species.
Virtual reality (today is the key to the past)
• Imagine traversing the habitats of the Molteno Biome at the time they
thrived. Walk those woodlands and riverine forests of the Late Triassic
as if you were walking their equivalents today. Feel their ecology and
biodiversity, conceive the interaction between plant, insect and verte-
brate species. Live those communities comprising biological species just
as today.
• Taxonomy and Late Triassic floristics will mirror reality more closely as
the virtual reality landscape comes into more vivid focus. Past reality
and present study merge in harmony.
Optimal sampling
• Optimal sampling of a formation will increase the chances of making
optimal taxonomic decisions.
• The more intensive/extensive the sampling of a fossiliferous formation,
the higher the proportion of correct decisions— all else being equal.
• When in doubt, increase the collection.
Curation & the palaeodeme approach
• The palaeodeme (population) approach— or something akin— is consid-
ered obligatory if the taxonomic goal is to approximate the original liv-
ing reality (pp. 22-25). Curation emphasising palaeodemes and tapho-
coenoses will maximise taxonomic reliability.
• Optimal curation involves the facilities necessary for the study of sever-
al palaeodemes simultaneously for comparative study. Moveable trays
(with many specimens showing variation within a palaeodeme) and a
closely adjacent, generously spacious, working surface on which to dis-
play an array of trays are pivotal.
Preparation & morphological data
• Dedicated preparation of specimens will optimise the proportion of cor-
rect taxonomic decisions.
• Cleanly exposing (chipping clear) a range of individuals within a
palaeodeme proves invaluable.
• Cuticular preparations and anatomical sections add to the inventory of
comparative data.
Objective & interpretive illustration
• Photographs (objective representations) and sketches (the authors' faith-
ful interpretation of morphology) of the best individuals covering a
palaeodeme are worth more than the proverbial plethora of words.
Taxonomic guidelines
c/tRELITZIA 15 (2003)
49
The lower cut-off
• It is in the nature of palaeobotany, no matter how small or large the col-
lection, that the rarer taxa and/or organs are more likely to be represent-
ed by poorer quality specimens,
• Where biodiversity is a core theme— as in this work— this innate prob-
lem has to be faced more emphatically than otherwise. Exacerbating the
problem is that in richer fossil floras (or insect faunas) a high proportion
of species appear both very rarely and very infrequently (pp. 20-25).
The more diverse the flora, the longer this tail.
• The lower cut-off in the quality of individuals accounted for in erecting
new taxa is necessarily lowered for rarer taxa.
• Specimens below the lower cut-off are those that are simply too inade-
quate for identity.
• Adequacy— an elusive measure— depends on the ability to see, sketch
or photograph definable diagnostic features on which the taxon can be
based (subjectivity is unavoidable).
The cone of uncertainty
• Uncertain decisions permeate palaeobotany; they are unavoidable and
are the very fabric of our research.
• All taxonomic decisions— including affiliations— are hypotheses (con-
cepts), some more sound than others.
• Such decisions range from near certain to distinctly uncertain, but they
cannot be avoided (assuming the material at hand falls above the nomi-
nated lower cut-off).
• Certainty increases with more material.
• Uncertainty increases with less material.
Controversial decisions (unanimity, mutuality, veto)
• The further down the cone of certainty-uncertainty, the less likely un-
animity will be reached.
• A veto rule has been applied in cases where a split vote has occurred
between ourselves (HMA and JMA) on a controversial decision.
• After due debate, each author secretly records his/her percentage vote
for a particular yes-no decision. (The votes ‘for’ may turn out to be as
decisive as 70-30% or as unsure as 49-51%). The average of the votes
of the two parties will be the decider. Further lobbying of viewpoints
may intervene before settlement.
Candidly documenting (and scoring) particularly uncertain decisions
(divergent votes) in a manuscript might prove valuably revealing,
e.g. JMA 70-30% for two particular species being congeneric;
HMA 49-51% against the two species being congeneric.
The balance of probability
• In the conventional ‘Criminal Justice System’, the particularly rigorous
guideline of ‘proof beyond reasonable doubt’ is employed: a person is
‘innocent unless proved guilty’. Even so, miscarriages of justice do
occur.
• In palaeobotany, were this rigorous guideline introduced, very few tax-
onomic decisions — at any level from species to class— would ever be
taken. We would emerge with a soup of words, but no taxa.
• The recent Truth and Reconciliation Commission in South Africa adopt-
ed the far less rigorous ‘balance of probability’ measure in assessing the
truth of a person’s testimony.
• This ‘balance of probability’ is the guideline underscoring much of our
palaeobotanical taxonomy.
4. MORPHOLOGICAL TERMINOLOGY
In the four-page spread that follows we present an illustrated key with
the aim of capturing all critical or specialised terminology used here in
describing the gymnospermous reproductive structures of the Molteno.
While there is no absolute consensus on usage in the more recent literature,
we have endeavoured to stay with the majority. We have therefore shifted
from our Prodromus of South African fossil floras (And. & And. 1985) in
which we chose to adopt the terms— such as fertiliger and polysperm —
introduced by Meyen (1987) specifically for gymnosperms.
No attempt has been made to include all Molteno fruiting genera com-
prehensively, as long as the key comparative morphology and terms are
covered. And for certain groups, to promote further clarity, non-Molteno
material is added. Pen sketches of the latter are included in a box with geo-
graphic and stratigraphic sources indicated.
The key serves also to summarise our interpretations of analogous and
homologous organs.
Analogous-, organs or parts having the same function but different evolu-
tionary origin, e.g. paddle of whale and fin of fish; convergent evolution.
Homologous', organs or parts having the same evolutionary origin, but dif-
ferent functions, e.g. paddle of whale and wing of bat; divergent evolution.
Ontogenetic variation (series, developmental stages)
Analysis of ontogenetic level — immature to senescent— at fossilisa-
tion can reveal interesting information on a range of related issues: taphon-
omy, affiliation, seasonality, diversity.
Cone buds— not found in the Molteno.
Immature (or abortive) cones — very rarely encountered in the Molteno, e.g.
Dordrechtites, 1 specimen (p. 64, tf. 9) from Lutherskop, and Hamshawvia,
a single strobilus from Aasvoelberg found attached to a short shoot with
Sphenobaiera leaves (pp. 210, 229).
Mature cones— with scales, bracts, seeds and pollen sacs (with in situ
pollen) intact; a small proportion of Molteno material, e.g. Rissikianthus
concavus, a fine palaeodeme of shed cones with in situ microsporangia
(p. 109) from Peninsula, and Kannaskoppia vincularis, a unique palaeo-
deme of strobili including a good many found attached to fragments of
shoot with or without Kannaskoppifolia leaves (pp. 288, 289).
Senescent cones— with scales, bracts, seeds, pollen sacs and pollen shed or
dehisced to lesser or greater degree; the great majority of Molteno repro-
ductive material falls in this category, e.g. Dordrechtites (pp. 60-69), fre-
quent and abundant in the Molteno, but almost exclusively found as
dehisced ovuliferous scales, and Kannaskoppianthus (pp. 290-293), with
virtually all strobili found detached and the great bulk having shed their
pollen sacs.
Strobilus
The term is used here generally for all gymnospermous male and
female reproductive organs, whether bearing compact or distantly spaced
sporophylls.
We use the following terms for different basic types of strobili:
Cone\ radially symmetrical compact strobili with imbricate or adjoining
scales— as for most female and male pinopsids (e.g. Telemachus,
Odyssianthus).
Pinnate or bipinnate ovulate ‘paniculate ' lamina, dorsiventral, bilaterally
symmetrical, laminate strobili, reminiscent of fertile fern fronds— as for
certain female ginkgoopsids (e.g. Umkomasia , Kannaskoppia).
Lax infructescense: ‘catkin’-like strobili with irregularly helical arrange-
ment of microsporangial heads — as for certain male ginkgoopsids (e.g.
Pteruchus, Stachyopitus).
Ottokariopsida (glossopterids)
On both the female and male spreads, we include a panel showing a
range of glossopterid reproductive structures, though the class of plants
remains unknown in the Molteno.
As recognised by various authors (White 1986; Retallack & Dilcher
1988; Anderson et al. 1999a), the glossopterids, absolutely dominant
through the Gondwana Permian and heavily decimated at the end-Permian
extinction, appear to have been a veritable nursery for new orders of gym-
nosperm (and possibly stem angiosperms) that arose during the extraordi-
nary radiation of new life through the Triassic.
In discussing the phylogeny and classification of the Molteno genera,
therefore, we make regular reference back to their possible ancestral glos-
sopterid lineages.
Taxonomic guidelines
50
c/tRELITZIA 15 (2003)
MORPHOLOGICAL TERMINOLOGY: female strobili
slender
pendent axis
P1NOPS1DA
Strobilus
radially-symmetrical
compact cone,
with helically attached
bract/scale complexes
J .stalk
Strobilus
scale 5-lobed, bearing pair
of adaxial seeds.
Megasporophyll
fascicle of 3-4 parallel imbricate
ovuliferous scales, dehiscing
individually from short stout stalk
Ovuliferous scale
I um-ovulate T-shaped scale, with
linear proximal & distal arms
xl
Dordreelitites
INCERTAE
Strobilus
pinnate “catkin”
/ Megasporophyll
/ sessile, ellipsoidal aggregate
-d of linear-lanceolate seeds
secondary
axis
Nataligma
Compound strobilus
gracile, leafless stem
with distant strobilus
whorls
Strobilus
pedunculate, ovoid
bearing several
sporopltyll whorls;
distal lamina '’fused to
form a closed cone.
GNETOPSIDA
Dispersed seed
(parent strobilus &
megasporophyll unknown)
leafy-wing with
parallel to forking
& anastomosing
venation
auriculate,
longitudinally
grooved seed -
Hirsutum
dliloi tides Megasporophyll
(fertiliger)
= bracuscale complex
of pinopsida
all x I (unless indicated)
Meyen ( 1987) terms in brackets
Morphological terminology
d^TRELITZIA 15 (2003)
5)
MORPHOLOGICAL TERMINOLOGY: female strobili
Strobilus
lax, bipinnate ovulate
“panicle”
Megasporophyil
pinnulate, with
2-3pairs of ovulate
cupules
MSi1
pedicel/ jl-2— axis
megasporophyil
Strobilus
radially symmetrical,
cylindrical,
compact cone
_ with helically arranged
j sporophylls
, Megasporophyil
\peltate
ovuliferous
disc
Strobilus
radially
symmetrical,
cylindrical,
semi-compact cone
with helically
arranged
sporophylls
Megasporophyil
palmate, recurved
ovuliferous
head
Peltaspernuim
Matatia
Fredl'india
peduncle —
"Cvnoecium
bilaterally
symmetrical
megasporophyil p
\ /TiX
cross-section
of gynoecium
BENNETTITOPSIDA
Megasporophyil (highly reduced)
biteemic ovules (consisting of an
ovide and its cupule)
interseminal scies
Morphological terminology
52
d^TRELITZIA 15 (2003)
MORPHOLOGICAL TERMINOLOGY: male strobili
CYCADOPSIDA
P1NOPSIDA
Microsporophyll
bilaterally symmetrical, simple scale,
sporophyll stalk, terete (circular in transverse
section, cylindrical and usually tapering),
distal lamina, heel (keel), inner surface,
external surface, peltate
episporangiate — adaxial
hvposporangiate — abaxial
peri sporangt ate — circum -axial
lateral
adaxial cluster
extant conifers: all pollen sacs develope
on abaxial surface
Microsporangium (pollen sac)
tfs 12-14 from Meyen (1987),
after Grauvogel-Stamm (1 978):
Gres-a-Voltzia, France, Triassic
Lutanthus Lutantlius Lutanthus Rissikianthus
robustus ornatus hemidiscus
Morphological terminology
^I/tRELITZIA 15 (2003)
53
MORPHOLOGICAL TERMINOLOGY: male strobili
GINKGOOPSIDA
nedicel
Strobilus
lax 'catkin', microsporophylls in
irregular helical arrangement
Strobilus
as for Slachyopilys
Microsporophyll
pedunculate, single or
aired laminate heads
'earing numerous
Dealing numerou
abaxial pendant
pollen sacs
Strobilus
lax, irregularly
branched
infructescence
Microporophyll
slender, pinnate,
sporangiate lamina;
pollen sacs in
radial, fascicled
clusters
secondary
axis (limb
micropsporophyll
microsporangium
Strobilus
bifurcate, pinnate,
microsporangiate lamina
Microsporophyll
single spathulate laminae
with dehiscent distal rim
subtending 5 pendulous
pollen sacs and protective
operculum
Kamuiskoppiu
BENNETT1TOPSIDA
W. spectabilis
“Flower” (bisporangiate cone)
Microsporophyll
(= pinnate frond)
arranged in a single whorl,
shaped like orange segments
Synangium (bean shaped)
contains 8-20 tubular
Cycadeoideu
sterile bracts
(helically arranged)
Williamsoniella coronata
monosporangiate cone (primitive condition) = dioecious
biosporangiate cone (derived) = monoecious
tfs 10-14 from Stewart &
Roth well (1993)
tf. 1 5 from Meyen ( 1 987 )
Euramerica,
Jurassic-Cretaceous
Leguminanthus
Microsporophyll — longitudinally folded toward
(presumed) abaxial surface
Pollen sacs — on inner surface, in small clusters of 3 or
4, arranged in irregular linear orientation
Weltrichia
Cycadolepis
Pen sketches (pp 52,53) are of Molteno
specimens unless otherwise indicated
Morphological terminology
54
d^TRELITZIA 15 (2003)
Tab. 30. CLASSIFIED LIST OF MOLTENO GYMNOSPERMS
DIVISION
CLASS
ORDER
FAMILY
Genus & species
PINOPHYTA A.Cronquist, A.Takhtajan & W.Zimmermann 1966
PINOPSIDA S.V.Meyen 1984
DORDRECHHTAI.ES order nov.
DORDRECHTITACEAE fam. nov.
Dordrechtites elongatus H.M.Ani
” cetipan’us sp. nov.
” mazocirrus sp. nov,
VOLTZIALES order nov.
FAMILIES INCERTAE SEDIS (2 families)
” omatus gen. et sp. nov.
” robustus gen. et sp. nov.
VOLTZIACEAE R.Florin 1951
• Telemachus elongatiis H.M.And. 1978
” grandis sp. nov.
” serribractus sp. nov.
” brachybractus sp. nov.
” dubibractus sp. nov.
” acutisquamus sp. nov.
Clariphyllum — 1 foliage
PINALES S.V.Meyen 1984
PODOCARPACEAE S.Endlicher 1847
Rissikistrobus plenus gen. et sp. nov.
” semireductus gen. et sp. nov.
” reductus gen. et sp. nov.
Rissikianthus concavus gen. et sp. no
linearis gen. et sp. nov.
” townrowii gen. et sp. nov.
” convectus gen. et sp. nov.
Rissikia — 2 foliage species
FAMILY INCERTAE SEDIS
Pagiophyllum — 1 foliage species
ORDER INCERTAE SEDIS
FAMILY INCERTAE SEDIS
ORDER INCERTAE SEDIS
FAMILY INCERTAE SEDIS
Helvetianthus tintinnabulum gen. et sp.
CYCADOPSIDA P.D.W.Bamard & A.G.Long 1975
CYCADALES A.Engler 1892
FAMILY INCERTAE SEDIS
Androstrobus peninsiformis sp. n<
” kraaiovalis sp. nov.
Pseudoctenis — 9 foliage species .
FAMILIES INCERTAE SEDIS (3 families)
J eanjacquesia — 3 foliage species
Clenis — 2 foliage species
Moltenia — 4 foliage species
GINKGOOPSIDA S.V.Meyen 1984
PELTASPERMALES S.V.Meyen 1984(?)
PELTASPERMACEAE H.H.Thomas ex T.M. Harris 1937
Peltaspermum thomasii T.M.Harris
monodiscum sp. nov.
tridiscum sp. nov.
turbanatum sp. nov.
quindiscum sp. nov.
Scytophyllum — 1 foliage species
MATATIELLALES order nov.
MATATIELLACEAE fam. nov.
Matatiella rosetta gen. et sp. nov.
” hemirosetta gen. et sp. nov.
” sessilis gen. et sp. nov.
” reducta gen. et sp. nov.
Kurtziana — 16 foliage spec
FAMILY INCERTAE SEDIS
Switzianthus moriformis ge
crispiformis gen. et sp. nov.
Dejerseya — 1 foliage spec!
GINKGOALES A.Engler 1897
AVATIACEAE fam. nov.
Avatia bifurcata gen. et sp. nov.
Eosteria eosteranlhus gen. et sp. nov
telemanthus gen. et sp. nov.
Ginkgoites — 6 foliage spec
FAMILY INCERTAE SEDIS
Classified list
^/tRELITZIA 15 (2003)
55
HAMSHAWVIALES order nov.
HAMSHAWVIACEAE fam. nov.
Hamshawvia baccata gen. et sp. nov.
” longipedunculata gen. et sp. nov.
” octosemina gen. et sp. nov.
” linisemina gen. et sp. nov.
Stachyopitys matatilongus sp. nov.
” matatiramus sp. nov.
” gypsianthus sp. nov.
” lacrisporangia sp. nov.
” rotundisporangia sp. nov.
i ” maziramus sp. nov.
Sphenobaiera — 9 foliage species
UMKOMASIALES order nov.
UMKOMASIACEAE S.V.Meyen 1984
Umkomasia macleanii H.H. Thomas 1933
” bracteolata sp. nov.
; ” quadripartita sp. nov.
” decussata sp. nov.
” monopartita sp. nov.
gracilliaxis sp. nov.
\ i ” capulata sp. nov. .
grandis sp. nov.
Pteruchus africanus H.H.Thomas 1933
matatimajor sp. nov.
” helvetigracilis sp. nov.
Dicroidium — 19 foliage species
Fanerotheca papilioformis sp. nov.
1 ” waldeckiformis sp. nov.
” cruciformis sp. nov.
” elandifortnis sp. nov.
PETR1ELLALES T.N.Taylor et al. 1994
KANNASKOPPIACEAE fam nov
Kannaskoppia vincularis gen. et sp. nov.
Kannaskoppianthus lutinumerus gen. et sp. nov.
matatiparvus gen. et sp. nov.
irregularis gen. et sp. nov.
telemagnus gen. et sp. nov.
Kannaskoppifolia — 10 foliage species
ORDER INCERTAE SEDIS
FAMILY INCERTAE SEDIS
Cetifructus bilateralis gen. et sp. nov.
CLASS INCERTAE SEDIS
ALEXIALES order nov.
ALEXIACEAE fam. nov.
Alexia urceolus gen. et sp. nov.
CLASS INCERTAE SEDIS
HLATIMBIALES order nov.
HLATIMBIACEAE fam. nov.
Hlatimbia tommacleanii gen. et sp. nov.
Batiopteris — 5 foliage species
CLASS INCERTAE SEDIS
ORDER INCERTAE SEDIS
FAMILY INCERTAE SEDIS
Hystricia perplexa gen. sp. nov.
CLASS INCERTAE SEDIS
ORDER INCERTAE SEDIS
FAMILY INCERTAE SEDIS
Saportaea — 1 foliage species
CLASS INCERTAE SEDIS
ORDER INCERTAE SEDIS
FAMILY INCERTAE SEDIS
Linguifolium — 1 foliage species
BENNETTITOPSIDA
FREDI.INDIALES order nov.
FREDLINDIACEAE fam. nov.
Fredlindia fontifructus gen. et sp. nov.
Weltrichia regalis sp. nov.
helvetirara sp. nov.
Cycadolepis rexiplumea sp. nov.
Leguminanthus leopardus sp. nov.
Halleyoctenis — 3 foliage species
PENTOXYLALES R.K.E.Pilger & H.Melchior 1954
LINDTHECACEAE fam. nov.
Lindtheca hackysackia gen. et sp. nov.
Taeniopteris — 8 foliage species
GNETOPSIDA H.G.A.Engler 1854
NATALIGMALES order nov.
NATALIGMACEAE fam. nov.
Nataligma dutoitii gen. et sp. nov.
Gontriglossa — 1 foliage species
ORDER INCERTAE SEDIS
FAMILIES INCERTAE SEDIS (2 families)
Graciliglossa — 1 foliage species
Cetiglossa — 1 foliage species
FRAXINOPSIALES order nov.
FRAXINOPSIACEAE fam. nov.
Fraxinopsis andium (J.Frenguelli 1941) comb. nov.
" auriculata sp. nov.
” comicordis sp. nov.
Yabeiella — 2 foliage species
Jungites — 2 foliage species
Classified list
56
d/TRELITZIA 15 (2003)
6. ON THE PHYLOGENY OF THE PINOPSIDA
With particular reference to the male cone genera and, more specifi-
cally, those from the Molteno.
State of knowledge
The phylogeny and classification of the genera of fossil coniferopsids
remain largely unresolved. Recent general texts reflect considerable differ-
ences in viewpoint (Meyen 1987; Stewart & Rothwell 1993; Taylor &
Taylor 1993). We adopt the order Voltziales in the sense of Stewart &
Rothwell (1993) as a group ‘transitional' between the Cordaitales and
Coniferales. and the Voltziaceae as a group intermediate between the
Utrechtiaceae and modem conifers. In particular we conceive the family
Voltziaceae as detailed in And. & And. (1989).
Male coniferous cones, aside from those found in the Molteno, are
entirely unknown in collections from elsewhere around the Gondwana
Triassic. Similar levels of infrequency (productive sites) and rarity (indi-
viduals per site) appear to be the pattern throughout the fossil record glob-
ally. They have added very little to our fragmentary knowledge of the phy-
logeny of the Pinopsida.
While the Molteno cones contribute substantially to the sum total
material available for study, they do not help clarify the lineages and rela-
tionships within the group. What they do suggest is that pinopsid evolution
during the Triassic may have been a good deal more complex than reflect-
ed in Tab. 31 opposite.
Ottokariopsida (glossoptedds)
As reflected in And. & And. (1985, pp. 106-145) and widespread else-
where in the literature, the Ottokariopsida and Cordaitanthales— apparent
stem group to all later pinopsids— have much in common. Their relation-
ship is certainly not resolved: did they share some common ancestor, did
the ottokariopsids arise from the Cordaitanthales, should they be grouped
more closely than shown here? We do not aim to explore these possibilities
further beyond commenting on the enigmatic options suggested by the
male conifer cones of the Molteno.
Did the southern conifers, or some clades thereof, evolve from the
glossopterids as suggested by White (1986, pp. 122-125)? Fredianthus,
Lutanthus and Odyssianthus, the Molteno male cones placed here within
the Voltziales, show certain features unlike those in the Laurasian
Carboniferous to Triassic voltzialean cones, yet shared by the glossopterid
family Lidgettoniaceae (And. & And. 1985, pp. 133-136; White 1986, pp.
118-121). These glossopterid features, appearing variously in the Molteno
cones, include the double row of microsporangia and the generally leafy
aspect to the scales — with most species characterised by distinct wings
(astride a midrib) along the scale axis and by the absence of a heel to the
distal lamina (i.e. being nonpeltate).
Cordaitanthales
Lea Grauvogel-Stamm (pers. comm., Pretoria, 15.11.1999) feels that
Fredianthus can be compared to the genus Cladostrobus (Maheshwari &
Meyen 1975; Meyen 1987), a cone associated with the leaf Rufloria and
found widespread in the Upper Permian of Siberia and Mongolia (see tf.
25, p. 59). Cladostrobus is a far smaller cone (over 60 mm long and ca 15
mm in diameter). The microsporophylls, consisting of a gracile terete stalk
and rhomboid distal lamina, bear groups of up to eight (oval to oval-ellip-
tical) microsporangia. Neither the degree of clustering, if any, nor the
nature of the attachment of the pollen sacs, however, is clear. Meyen (1987)
reconstructs and describes the microsporophylls as perisporangiate (sacs all
around the stalk) rather than hyposporangiate (adaxial), with no suggestion
of clustering.
It is possible that Fredianthus , in the Molteno, represents a distinct,
surviving cordaitanthalean lineage rather than falling in the mainstream of
the Voltziales. Short of further information, we follow the voltzialean
option.
Voltziales
Voltziaceaen foliage and female cones are very characteristic compo-
nents of global floras from the Upper Permian to Middle Jurassic (And. &
And. 1989, pp. 420^123), yet the male cones barely make an appearance.
As far as we are aware, the only previously described polleniferous cones
reasonably established as belonging to the family are the group of genera
Sertostrobus, Darneya and Willsiostrobus deriving from the lower Middle
Triassic Voltzia Sandstone and approximate time equivalents elsewhere in
Europe and beyond in Laurasia (Grauvogel-Stamm 1978; Taylor 1988;
Grauvogel-Stamm & Galtier 1998). Well preserved Willsiostrobus speci-
mens have also been described from late Lower Triassic beds in N. China
(Wang & Wang 1990). A further record is that of Krassilov (1982): D.
angusta and W. latisaccus from the Lower Cretaceous of Mongolia.
The three genera of putative voltzialean male cones,
Fredianthus , Lutanthus and Odyssianthus , described here from the
Molteno, add, as previously noted, significantly to the spectrum of taxa and
morphology at hand.
Although largely distinct from the European voltziaceaen male cones,
the three Molteno forms are all included provisionally in the order
Voltziales: Fredianthus and Lutanthus, without any foliage or megasporan-
giate affiliates, in two unspecified families; Odyssianthus, with well-estab-
lished foliage ( Heidiphyllum ) and female-cone ( Telemachus ) affiliates, in
the Voltziaceae. Both Heidiphyllum and Telemachus are undisputed mem-
bers of the family Voltiaceae as currently constituted.
Odyssianthus with its two rows of latero-abaxial, clustered, sessile
pollen sacs, Fredianthus with its single row of abaxial, clustered, near-ses-
sile sacs, and Lutanthus covering a wide spread of morphological types,
hint at family- and order-level diversity in the Late Triassic pinopsids
markedly richer than reflected in our present classification based on ovu-
late material alone.
The Voltzia Sandstone genera likewise show considerable differences
from one another, suggesting, already in the early Middle Triassic, a diver-
sity amongst the nonpinalean pinopsids not yet fully recognised. While
Darneya and Sertostrobus bear adaxial sporangia along the stalk,
Willsiostrobus sports pollen sacs that are abaxially attached to the heel of
the distal lamina. Grauvogel-Stamm & Galtier (1998), moreover, interpret
Darneya as bearing a complex microsporophyll— with fused scale and
bract— and propose the presence of two strongly divergent lineages of
conifer in their material. Perhaps the three French genera represent three
major lineages, only one of which is voltzialean.
Aside from the Molteno material, male cones of the order remain
unknown from the Gondwana Triassic— although Heidiphyllum foliage is
very widespread and often a dominant component of assemblages.
Pinales
Both Lebachia, included in the Utrechtiaceae (Lebachiaceae, Walchi-
aceae), and Willsiostrobus (Middle Triassic, Europe) in the Voltziaceae,
show horizontally alligned microsporangia attached to the distal lamina of
a strongly peltate scale. These are features almost ubiquitously characteris-
ing the Pinales, the diverse crown group of Pinopsida. Rissikianthus, from
the Molteno, likewise shares these features. Did such characters arise inde-
pendently several times within the stem Pinopsida; is some definite reshuf-
fling of taxa within the Carboniferous to Triassic families indicated; are the
families quite wrongly conceived at present? The reality of pinopsid phy-
logeny is as yet elusive. A quantum jump in the available sample of male
cones will, no doubt, contribute to the resolution sought.
Primitive & derived characters
If it is true, as generally supposed, that the Cordaitales are the stem
group leading to all later pinopsids (and possibly the ottokariopsids), then
it is amongst this group back in the Early to Middle Carboniferous that we
must seek their primitive characters. Unfortunately male cones of this crit-
ical vintage and order appear unknown. We move then directly to the
Voltziales and Ottokariales of the latest Carboniferous and Permian (see
Tab. 31).
At least three markedly different kinds of cone already existed: in
Dvinostrobus, Lebachia and Lidgettonia. They differ in virtually every
diagnostic feature characterising male cones of these late Palaeozoic
orders. With this morphological spectrum as a starting point, all features
defining these genera are witnessed as equally primitive and all features
newly evident in Mesozoic pinopsid males as derived.
Phylogeny of Pinopsida
c/tRELITZIA 15 (2003)
57
Phylogeny of Pinopsida
58
d>^TRELITZIA 15 (2003)
MOLTENO PINOPSIDA: male cones
Tab. 32. Diagnostic features of the four voltzialean & pinalean genera
W ,
Pinales
Fredianthus
Luthanthus
Odyssianthus
Rissikianthus
Strobilus
Size
to 220 mm in length
30-40 mm in length
70 mm in length
10-18 mm in length
Gyres
60
13-24
23
7-10
of 25-30 microsporophylls
of 10-20 microsporophylls
of 16-20 microsporophylls
of 6-12 microsporophylls
Axis
moderately flexed
strongly flexed to erect
strongly flexed to base
strongly flexed to base
Microsporophyll
Stalk
sigmoidal
curving upward distally
strongly upward-curving
straight
Lamina
narrowly ovate, leafy
broad, leafy
broadly ovate, leafy
triangular, woody
multilobed/dentate.
entire to multi-ribbed.
finely crenulated,
entire,
without heel
without heel
with moderate heel
with strong heel
Microsporangia
single median row.
1 median or 2 lateral rows.
in 2 lateral rows,
a single pair,
5 clusters along the row,
3—4 singles or 2 clusters,
4 clusters per row,
1 single per side,
45 sacs per scale,
6-8 sacs per scale,
22-24 sacs per scale,
2 sacs per scale,
abaxial (along stalk)
adaxial to abaxial (along stalk)
latero-abaxiai (along stalk)
abaxial (from heel)
Microsporangium
Shape
elliptical
obovate to rhomboidal
irregularly rhomboidal
rotund
Dehiscence
clear longitudinal line
usually with clear longitudinal
clear longitudinal line &
longitudinally bilobed
line & apical “micropyle"
apical "micropyle"
(no obvious dehiscence)
All numbers (with a few exceptions) are approximate
Phylogeny of Pinopsida
t/tRELITZIA 15 (2003)
59
EXTANT
(Global)
'The 2 pollen sacs & disaccate pollen
o{“l(issikia" are found with regularity
in the Podocarpaceae & Pinacea’e
only” — Townrow 1 967 (seep. 105)
PINACEAE ' ^rorn Foster & Gifford (1974)
Pinus
, 2
Class PINOPSIDA: male cones
PODOCARPACEAE
Podocarpus tf. 2 front Leistner (1966)
Abaxial
ARAUCARIACEAE
Araucaria
tf. 8 from
Marsh ( 1 966)
CUPRESSACEAE
Cupressus
TAXOD1ACEAE
Cunningham ia / j
xii
tfs 3-7 from Page (1990)
tf, 9 from
Foster & Gifford (1974)
MESOZOIC
(Laurasia)
tfs 10-15 from
Grauvogel-Stamm (1978)
Gres a Voltzia, France, Triassic
Darneya
CHEIROLEPID1ALES
CHEIROLEP1D1ACEAE
Sertostrobus
VOLTZIALES
VOLTZIACEAE
microsporophyll
18
Classopollis
tfs 1 6-1 8 redrawn
from Watson (1982),
Jurassic-Cretaceous,
Western Europe
LATER PALAEOZOIC
Lidgettonia
OTTOKARIOPSIDA
LIDGETTONIACEAE
(Gondwana Permian>
tfs 19-21 from
And. & And. (1985)
Sq name l la
CORDA1TANTHALES
RUFLORIACEAE
Cladostrobus lutugimi
Siberia, Penman
VOLTZIALES
(Laurasia)
LEBACHIACEAE
(=Utrechtiaceae, =Walchiaceae)
26
EMPORIACEAE
(the most primitive
voltzialean family)
Lpinformis Lebachia
Western Europe
Lower Permian
Phylogeny of Pinopsida
60
c/tRELITZI a 15 (2003)
PINOPSIDA S.V.Meyen 1984
DORDRECHTITALES J.M.And. & H.M.And., ord. nov.
DORDRECHTITACEAE J.M.And. & H.M.And., fam. nov.
Dordrechtites H.M.And. 1978
Type species
Dordrechtites elongatus H.M.And. 1978.
Generic diagnosis
A pinopsid female cone bearing fascicles of gracile, T-shaped, ovulate
scales that detach readily from short stout pedicels.
Generic characters
Attachment : unknown.
Strobilus: simple, compact to lax cone, broadly linear, medium to large (to
ca 150 X 22 mm); axis slender, curved, pendent; scale clusters subop-
posite, subdecussate.
Megasporophyll: in fascicles of 3 or 4 parallel to partly overlapping ovulif-
erous scales, readily detaching from short stout pedicels.
Ovuliferous scale ; T-shaped (30 X 10 mm); sterile arms of T roughly
equal, gracile, linear, dorsiventraily flattened; proximal arm strongly
arcuate, tapering to square apex at attachment; distal arm mildly arcu-
ate, tapering to finely acute tip; ovuliferous trunk of T relatively short,
robust, triangular in side view, curving proximally, dorsiventraily flat-
tened, broadly ovate to obovate in plan view, with strong ventral keel,
broad lateral laminar wings and mucronate tip.
Ovule/seed: adaxial, enclosed dorsally within winged ovuliferous trunk
and never found detached, narrowly ovate in plan, dorsiventral.
Cuticular features', see text below.
Etymology
Dordrechtites— after the town Dordrecht situated near the type locality.
Global range: 4 spp., Gondwana, Tr. (LAD-CRN).
First: Dordrechtites sp. (Holmes, pers. comm.); Dubbo district, N.S.W.,
Australia.
Last: Dordrechtites sp. (HMA, this vol., p. 61 ); Aqua de la Pena, Ischigua-
lasto, Argentina.
Gondwana Triassic occurrence
SAm— N. Argentina, 1 loc. (>6 indivs).
SAf— Karoo Basin, 18 TCs (>413 indivs).
Aus — Queensland and New South Wales, 2 Iocs (>120 indivs).
Molteno occurrence
Frequency (F): 17 TCs (of 100 sampled in Molteno).
Diversity (D): 3 species.
Abundance (A): 413 individuals, common to very rare.
Lut 311 Hei elo: >50 indivs in 50 man-hrs cleaving (common)
Aas 41 1 Dic/Sph:
>100 ’
” 512
” (occasional)
Aas 311 Hei elo:
>40 ’
” 140
(occasional)
Bir 1 1 1 Sph 2spp:
>50 ’
” 550
” (occasional)
Maz 211 Hei/Dic:
40 ’
” 85 ”
” (5 per 1 man-day) rare
Lit 111 Dic/Hei:
>50 ’
” 550 ”
” (>1 per 1 man-day) rare
Umk 1 1 1 Die 2spp
10 ’
" 400
( 1 per 4 man-days) very rare
Whereas intact or partially intact cones of Dordrechtites are extreme-
ly to vanishingly rare, the dehisced ovuliferous scales are, in a good many
TCs and especially on certain bedding planes, occasional to common.
Affiliated organs
Male cone: unknown.
Foliage: unknown (but see p. 62).
Classification & comparison
Suprageneric classification (Dordrechtitaceae/Dordrechtitales)
The cones and ovuliferous scales of Dordrechtites offer a pinopsid
impression, but the extraordinary mode of attachment of the scales in pedi-
cellate fascicles is unique (either as reconstructed here, or as interpreted in
Playford et al. 1982, for the Australian species— see p. 63). While it would
stretch comparisons unduly to consider Dordrechtites as falling either with-
in the Voltziales or the Pinales, it seems reasonable to include the genus in
the class Pinopsida as a new family (Dordrechtitaceae) and order
(Dordrechtitales). The Voltziales represent the most likely known sister
clade.
Intergeneric comparison (Gondwana Triassic)
Nothing comparable is known.
Dordrechtites
DORDRECHTITALES
TRELITZIA 15 (2003)
61
Reconstructions
The set of Grade 3-5 reconstructions (p. 60, tfs l^t) of the type
species, D. elongatus , is based largely on the selection of specimens (Grade
2 reconstructions, p. 64 [1-10]) from the reference palaeodeme and top
three sister palaeodemes — Aas 4 1 1 , B ir 1 1 1 , Aas 311 and Lut 3 1 1 — for the
species. In view of the morphological uniqueness of this genus, we elabo-
rate more fully than usual on the relative certainty of the reconstructions.
Mature cone (p. 60, tfs 1, 3)
The mature strobilus is fairly confidently portrayed as terminal, pen-
dulous and occurring singly at the end of arching branchlets. Its length is
based on the most substantial specimen from Aas 41 1 (PRE/F/21 762 a,b; p.
64, tf. 3), backed up by a bare axis of ca 150 mm from the Moolayember
Fm. of Australia (Playford et al. 1982, pi. 2, fig. 4), with the base appar-
ently preserved, but the apex missing. The pendulous nature is deduced
from the gracile, generally curving axis and the strongly curved proximal
end in the single available immature specimen (Lut 311, PRE/F/11439, p.
60, tf. 2a,b). The subopposite, subdecussate attachment of the scale clusters
is based on PRE/F/21762 a,b (p. 64, tf. 3) from Aas 411 and PRE/F/22509
(p. 64, tf. 4) from Aas 311.
Immature cone (p. 60, tf. 2a, b)
The single, partly intact cone from Lut 311 (PRE/F/1 1439; p. 64, tf. 9)
bears scales less than half normal size. The specimen could represent a dis-
tinct species, but since all further single or clustered scales from Lut 311 (p.
64. tfs 8, 10) are like those of D. elongatus from other TCs in size and form,
it is assumed to be an immature cone.
Ovuliferous scales (p. 60, tfs 4a-d)
The number of scales in the scale clusters on a mature intact cone
remains uncertain. Nowhere in the few partly intact cones at hand (p. 64,
tfs 2-4, 9, 10) is the number sure. It is evident that in the closed cone the
arms of the scales lie immediately adjacent to one another, while the
winged ovules overlap strongly. We favour clusters of four scales (only
slightly over three) as the norm and have drawn the reconstructions on this
basis.
Ovule & seed
While the great majority of dehisced scales are preserved in lateral
view, they are very occasionally seen straight on with the ovule in full or
partial dorsiventral aspect (p. 64, tfs 5-7, 10) allowing the interpretation as
shown. The ovuliferous part of the scale is clearly dorsiventrally flattened
and is markedly winged. In only one specimen (Aas 311; PRE/F/22515; p.
64. tf. 6) is the intact seed preserved.
Classification (elaborated)
Laurasia Triassic
The closest apparent comparison amongst Voltzialean genera is with
Borysthenia of Russia, Donets Basin, Upper Triassic (Stanislavsky 1976,
p. 77-81, pi. 43-47; And. & And. 1989, p. 423). Aside from the scales,
Dordrechtites is distinct in having clear pedicels from which the scale
fascicles detach.
Other ages
The seed of the extant Araucaria (Coniferales) is superficially similar
to the central winged seed/ovule of Dordrechtites — which is, however,
invariably found with the two linear projections.
Some similarity occurs with the winged seed Semenalatum paucum
(Dilcher et al. 1997) from the Early Permian of China and thought by them
to have coniferous affinities. This seed, however, is characterised by two
lateral wings and one limited median projection, and the point of attach-
ment is central where three additional, short, sterile scales occur.
In other plant divisions we have encountered two fossils that bear a
remote resemblance to the Dordrechtites scale. Douglas (1969) describes
an isolated angiosperm seed, Lappacarpus, with two appendages, but these
are very thin and only 2-5 mm long. Within the Lycophyta, Cantheliophorus
(Thomas & Brack-Hanes 1991) shows only a single extension, but this is
clearly leaf-like with a single midrib.
Gondwana Triassic occurrence (elaborated)
Dordrechtites, frequent and abundant in the Molteno — at least as
dehisced scales— remains very sparsely known elsewhere in the Gondwana
Triassic. Only three TCs yielding the genus are known outside of Africa.
South America
Definite Dordrechtites scales with Heidiphyllum leaves were recorded
on a bedding plane by HMA (unpublished notes, 23-09-1999) at Agua de
la Pena locality (Los Rastros Fm.) during the VII International Symposium
on Mesozoic Terrestrial Ecosystems field excursion to Ischigualasto,
Argentina. No material was collected.
Australia
A well-represented Dordrechtites palaeodeme, including two broken
cones (one being indistinctly preserved) and over 120 isolated scales, is
known from a single ‘locality' in the Lower Ladinian Moolayember Fm.,
Bowen Basin, Queensland (Playford et al. 1982; for further details see pp.
62, 63). This remains the only published occurrence of the genus outside
the Molteno. Keith Holmes (pers. comm.) has seen rare isolated scales at a
locality in the Dubbo district. New South Wales.
GONDWANA TRIASSIC, GEOSTRAT DISTRIBUTION
DORDRECHTITALES
Dordrechtites
62
d/TRELITZIA 15 (2003)
assemblages
(taphocoenosis)
Heidiphyllum
O Telemachus
+<j Dordrechtites
1-2 scales a
3-10 scales jj. §
(clusters) g
>10 scales = 2.
(clusters) $ -»
scales m ?
attached
man-hours
cleaving
Nav 111 Die odo
1 - -
2
Cal 211 Heielo
75 - -
- i - : - i -
2
Bir 211 Sph 2spp
3 - -
.
7
" 311 Hei/Sph
45 - 6
6 -
2
” 111 Sph 2spp
10 - 50
47 2 1
550
Dor 111 Heielo
85 - -
.
2
Gre 121 Hei elo
98 100 -
- i - 1 - i -
10
” 111 Sph pon
10 - -
_ | - i - !
5
Boe 111 Lep sto
7 - -
- ! - ! - -
8
’’ ” Dic/Hei
42 - -
.
8
’’ 112 Die cor
14 - 2
2 - - -
6
Cyp 111 Die era
24 5 -
-l-i - ; -
100
” 111 Hei elo
100 - -
- : -
2
Kan 112 Hei elo
98 10 -
- - : -
15
" 111 Ast spA
10 - -
- 1 - - 1 -
30
Tel 111 Hei elo
89 40 -
- i -
90
Vin 111 Die odo
28 - -
-
10
Ela 111 Dicoco
7 1 -
- ' - - -
10
Kra 311 Die odo
5 - -
- J - ; - -
13
Lut 111 Hei/Dic
50 2 -
- - - -
2
” 511 Heielo
80 1 -
. ; . - : -
2
" 411 Hei/Dic
50 12 -
- ; - : -1 -
2
" 311 Hei elo
99 - 50
47 1 1 1
50
Tin 121 Sph 2spp
4 - -
- -
5
" 111 Sch sp
10 - -
- ■ -
3
" 131 Hei/Ast
50 - -
- ! - - -
3
Kon 223 Die odo
1 - -
-i-l - [ -
7
” 211 Heielo
95 - -
- ; - ; - -
1
” 111 Die odo
7 - 1
i
34
” ” Equ sp
10 - -
-
1
" " Hei elo
84 2 18
9 5 4 -
4
Pen 321 Dic/Ris
1 1 -
- 1 -
35
” 211 Dic/Equ
2 - -
-|-1 - -
4
” 421 Die odo
4 - -
- ; - ; - -
7
” 431 Dic/Equ
5 - -
- i - - ■ -
1
" 311 Heielo
75 17 5
5 - - -
35
■■ 41 1 ” "
94 50 1
1 - - -
70
Klelll Neo car
1 - -
- ; - ; - -
4
” ’’ Hei elo
90 25 -
- : - - ; -
15
” ’’ Hei/Dic
49 - -
- ; - -
9
Kap 111 Dic/Ris
25 5 -
- i - - 1 -
65
Ela 112 Dic/Hei
30 - -
-1-1 - -
4
Nuw 211 Die 2spp
1 - 4
4 - - -
6
Win 111 Heielo
79 - 15
6 3 6 -
20
Qua 111 Die odo
20 - -
- ■ - : -
8
Maz 111 Die era
5 - -
- 1 - i - 1 -
30
" 211 Hei/Dic
32 - 40
38 2
85
Hla 213 Die elo
i -
- ; - -
60
Umk 111 Die 2spp
7 - 10
10
400
1 nj 211 Die dub
10 - -
.
2
San 111 Die era
5 - 15
15 - -; -
30
Mng 111 Die 2spp
6 - -
- 1 - 1 - 1 -
2
Qac 111 Hei/Dic
50 10 -
- : -
4
Mat 111 Die dub
4 12 -
-
65
Lit 111 Dic/Hei
23 - 50
46 4
550
Aas 611 Heielo
80 6 -
... -
3
” 111 ” "
77 12 -
- i -! - l -
40
„ 211 „ „
100 - 6
6 -
35
” 311 ” ”
99 - 40
37 1 1 1
140
” 411 Dic/Sph
1 - 100
67 15 15 3
512
” 511 Die elo
20 - -
.
2
Bam 111 Die dub
1 - -
-1-1 -| -
3
Total TCs
62 18 17
17 8 6 3
Total indivs
% 311 413
347 33 28 5
Tab. 33. Dordrechtites, Molteno occurrence
Abundance: Dordrechtites may be quite abundant at cer-
tain TCs and it is possible to collect numerous individuals.
In these cases, we record the number of individuals
(rounded off where ca 40 indivs or above) as curated for
the genus, but do not include further specimens which
occur on slabs in the remainder of the collection.
Evidence for affiliation of organs
Dordrechtites & Telemachus (mutually exclusive genera )
The distribution pattern of the two ovulate organs, Dordrechtites and
Telemachus, in the 100 Molteno assemblages is an intriguing one (Tab. 33).
Dordrechtites , a cone scale of still uncertain affinity, is known from 17
assemblages; Telemachus, the voltzialean coniferous cone affiliating with
the abundant leaf Heidiphyllum, is known from 18 assemblages. Both genera
are invariably found in beds yielding Heidiphyllum— known from 62
assemblages, often as the monodominant element— but their occurrence is
virtually mutually exclusive. What does this striking pattern suggest with
regard to depositional environment, taphonomy and affiliation?
On Heidiphyllum (two foliage genera or one?)
An option that has been entertained is that the genus Heidiphyllum
actually includes two quite distinct, though superficially similar, foliage
genera. [We have previously, in our study of the Molteno flora, separated
out Sphenobaiera insecta from Umk 1 1 1 (And. & And. 1989, p. 144). The
taxon looks very like Heidiphyllum in shape, size and venation, but is
undoubtedly a species of Sphenobaiera based on cuticle and the finer
details of venation.] After close examination of a good number of
Heidiphyllum palaeodemes from TCs including either Dordrechtites or
Telemachus, we found no sign of two foliage genera. A very characteristic
feature of Heidiphyllum, allowing its sure identification, is the clear pres-
ence of interveinal striae (And. & And. 1989, pis 249-263). This occurs in
all palaeodemes examined.
Dordrechtites as a pinopsid (the taphonomic filter)
If Dordrechtites is correctly classified as representing a new order in
the class Pinopsida, we might expect the foliage of the plant to appear
typically coniferalean. In view of the frequent and abundant occurrence
of Dordrechtites in Molteno strata one would also imagine the parent
plant to be a prominent element characterising certain communities in the
Molteno Biome. How is it that the foliage could have been totally filtered
out in the taphonomic/fossilisation process? The question remains unre-
solved.
The pinopsid-foliage enigma does not end with Dordrechtites. We
have identified five pinopsid male-cone genera in the Molteno, for three of
which, Fredianthus, Lutanthus and Helvetianthus, there occur no evident
foliage affiliates (Tab. 34, p. 75). And we recognise five female-cone gen-
era (only two with male-cone affiliates), three of which, Dordrechtites,
Gypsistrobus and Avistrobus, likewise have no evident foliage affiliates.
These other fruit genera, unlike Dordrechtites, are all extremely infrequent
and rare. Overall, then, there occur six pinopsid whole-plant genera repre-
sented by male or female cones, but whose foliage, apparently, has been
comprehensively filtered through the taphonomic net.
The particular difficulty in establishing affiliations between dispersed
pinopsid organs in a well-sampled formation is not unique to the Molteno.
The Voltzia Sandstones of the French Lower Triassic yield a rich pinopsid-
dominated flora with several distinctive female, male and foliage genera.
After careful curation, by localities (TCs) then taxa, most affiliations
remain a puzzle (Grauvogel-Stamm & Anderson J.M. 1980, unpublished
notes).
Dordrechtites, an allochthonous vagrant
There is a marked, though not exclusive, correlation between
Dordrechtites and more well-bedded, lower-flow-regime deposits (most
notably Bir 1 1 1 and Aas 41 1 ) on the one hand, and between Telemachus
and more poorly laminated, higher-flow-regime deposits (most notably Tel
111 and Kan 112) on the other.
The most likely resolution, as we expressed in Caimcross et at. ( 1995),
is as follows: ‘It is significant that Dordrechtites is a winged scale, appar-
ently adapted for wind (and water) dispersal, and that these scales are
invariably detached and usually scattered through a deposit. They almost
certainly represent communities growing some distance removed from the
site of deposition. Telemachus. on the other hand, is a 5-6 cm long woody
cone, often found intact or partially intact, although isolated scales are also
common. They appear mostly to have undergone little transportation and to
have been derived from the coniferous community closely adjacent to, or
within the site of deposition.’
Australia (Moolayember Fm .)
The depositional environment of the Moolayember Fm. is interpreted as
a fairly extensive, inland, fluviatile-lacustrine basin with some evidence of
intermittent brackish and ephemeral marine incursions (Playford et al.
1982). The Dordrechtites material derives from a single TC (‘locality’) and
consists of ‘at least 120 seeds attached to cupules, one broken cone ...., and
an indistinctly preserved broken cone'. We recognise, in addition, the ‘gym-
nosperm stem’ illustrated by Playford et al. ( 1982, pi. 2, f. 4) as the axis of
a Dordrechtites strobilus with numerous short pedicels.
Dordrechtites
DORDRECHTITALES
'"“'/trELITZIA 15 (2003)
63
The associated flora, as reflected in Playford et al. (1982), consists of
stems and foliage of a single species of horsetail, four genera and species
of fern, Dicroidium (two fragments), Linguifolium (a single fragment),
Ginkgoites (one individual), a few other ginkgophyte fragments, cfRissikia
(four leafy shoots) and some fair-sized fragments of leafless gymnosper-
mous stem. Gymnosperm fruit, aside from Dordrechtites, consist of
Umkomasia (a few individuals, with one fairly complete strobilus) and
Pieruchus (one fragment). The content and preservation of this assemblage,
with the intact Dordrechtites and Umkomasia strobili and the gymnosperm
stem fragments, suggests near autochthony and relatively rapid burial.
One of us (HMA, 1988) had the opportunity of studying the original
collection— housed in the Dept, of Geology & Mineralogy, University of
Queensland, Brisbane — and observed numerous Heidiphyllum leaves on
slabs from this locality. These leaves were neither described nor specifical-
ly illustrated in Playford et al. ( 1982), though two fragmentary specimens
do appear on the slab (pi. 6, f. 6) showing the partially intact Dordrechtites
dikeressa cone.
The Spring Creek TC, then, like many Molteno TCs, yields the
Dordrechtites/ Heidiphyllum rather than the Telemachus/ Heidiphyllum
association of taxa, but does not contribute towards resolving the
Dordrechtites affiliation dilemma. No other vegetative element in the flora
appears to be a likely affiliate of Dordrechtites.
South America (Los Rastros Fm.)
Further evidence of Dordrechtites found in association with
Heidiphyllum. was noted by HMA (23-09-1999) at the Aqua de la Pena
locality, Ischigualasto, where a bedding plane was found literally covered
with specimens of the two genera.
Intactness of cones
The overwhelming majority of Dordrechtites specimens are preserved
as isolated scales. They obviously separate readily from the stout pedicels
to which they are attached in fascicles of three or four. The proportion of
dispersed single scales to scale clusters or partial strobili in the deposits, is
far higher than is evident in the table— which reflects curated material only,
not counts on site. For those TCs with more numerous Dordrechtites
remains (e.g. Lut 311, Aas 311, Aas 411), the collection is strongly biased
towards scale clusters (from one or more cones) and cone fragments. These
are preferentially retained, while a great number of lesser preserved single
scales are discarded.
It is difficult to discern any revealing pattern between the occurrence
of scale clusters or cone fragments and habitat/TC type. Aas 411 (a lake
deposit) is the top fruit-yielding TC in the Molteno with 13 genera of ovu-
late fruit, some of which are notably intact, including three specimens of
Dordrechtites. Bir 111 (the other prolifically fossiliferous lake deposit)
with six ovulate cone genera has yielded almost exclusively dispersed
scales. Aas 3 1 1 and Lut 311, each with a single cone fragment, a couple of
scale clusters and numerous single scales, represent low-diversity
Heidiphyllum thickets. Kon 1 1 1 and Win 1 1 1, the additional two TCs with
relatively high proportions of scale clusters, likewise represent low-diver-
sity Heidiphyllum thickets.
Cuticles
Potential sample: Lit 1 1 1, ca 50 dispersed scales; Umk 111, 10 scales.
Macerated (this work): Lit 111,5 scales.
Preservation grade : Grade 2 (see JSM results below).
Diagnostic characters: cells oblong, pentagonal-hexagonal, walls gently
curved; nonpapillate; stoma orientation longitudinal, anomocytic;
other features absent.
Comment: in the absence of results from maceration, the Jed Scanning
Microscope (58001v) scan was attempted on two or three fragments.
The features sketched and described are based on this technique.
Significance:
Classification— In its cellular features (shape, walls) and the thinness
of its cuticle, Dordrechtites is more readily placed in the pinopsids than
other classes such as the ginkgoopsids or gnetopsids. The cuticle is similar
to that from the upper leaf surface of Heidiphyllum (p. 90), which has
anomocytic stomata and elongate cells. Rissikia (p. 112) cuticles also have
oblong cells, but the stomata are brachyparacytic and lappetate. It is cer-
tainly very unlike the cuticle of the pinopsid genus Pagiophyllum (p. 124)
with its regularly actinocytic subsidiary cells which are strongly cutinised
and lappetate. The cycadopsids cannot be ruled out, as some (e.g.
Pseudoctenis, p. 140) also have anomocytic stomata and elongate cells.
Other poorly known or unknown classes of plant may obviously enter the
picture.
Affiliations— The cuticle, in being similar to that of the leaf genus
Heidiphyllum. adds to the dilemma (p. 62) around Dordrechtites. in addi-
tion to Telemachus. being a candidate for affiliation with this leaf.
Adaptive radiation (Molteno diversity)
Though Dordrechtites is both frequent (17 TCs) and often relatively
abundant (>50 individuals per TC) in the Molteno, little diversity can be
readily discerned. In view of the extreme scarcity of intact strobili— all
appearing very alike— this assessment is based essentially on the ovulifer-
ous scales alone. The diagnostic features characterising the species lie in
the relative shape, robustness and size of the sterile arms and fertile trunk
of these scales (as preserved in lateral view).
The three Molteno species are based on the following TCs/reference
palaeodemes. Each derives from a distinct habitat and from a different
level within the stratigraphic sequence.
D. elongatus— Aas 41 1 Dic/Sph (Aasvoelberg), 100 indivs
Sphenobaiera closed woodland; Cycle 1 (Bamboesberg Member)
D. cetiparvus— Umk 111 Die 2spp (Umkomaas Valley), 10 indivs
Dicroidium riparian forest (mature); Cycle 2b (Indwe Member)
D. mazocirrus—Maz 211 Hei/Dic (Mazenod), 40 indivs
Dicroidium riparian forest (immature); Cycle 2c (Indwe Member)
sketched from Jed
Scanning Microscope
(5800 Lvj photo
D. elongation
DORDRECHTITALES
Dordrechtites
64
TRELITZI A 15 (2003)
Dordreclitites elongatus H.M.And. 1 978
Holotype
Specimen: BP/2/5283(C-Dt.II 365 in And. 1978); pi. 3(3).
Assemblage (TC): Bir 111 Sph 2spp; Birds River.
Presentation: single dehisced scale, without counterpart; impression in
thinly laminated, yellowish grey shale with very good cleavage.
Reference palaeodeme
Assemblage (TC): Aas 411 Dic/Sph; Aasvoelberg; pi. 1( 1-8).
Specimens: >100 individuals; 2 incomplete mature strobili (tfs 2, 3 adja-
cent), 1 incomplete immature strobilus, ca 15 scale clusters of >10 de-
hisced scales and ca 15 clusters of 3-10 scales; many isolated scales.
Presentation: impressions (3D moulds/casts) in thinly laminated, strongly
baked, yellowish grey shale with very good cleavage.
Sister palaeodemes— 15 (best three listed)
Bir 111 Sph 2spp: >50 indivs (1 good scale cluster).
Lut 311 Hei/Dic: >50 indivs (1 strobilus).
Aas 311 Hei elo: >40 indivs (1 strobilus).
Specific diagnosis
A Dordrechtites species bearing relatively large scales with long,
gracile, roughly equal arms and gently arching fertile trunk.
Specific characters
Ovuliferous scale: relatively large; sterile arms gracile, approximately
equal (15 mm long); fertile trunk acutely conical, arching slightly
proximally.
Etymology
elongatus (Lat.)— with reference to the long arms of the scale.
Comments & comparison
This is by far the most common species of Dordrechtites, with the
scales being remarkably constant in size and shape through the 17 differ-
ent TCs. It is distinct in being larger and with longer arms than both D.
cetipan’us which is considerably smaller, and D. mazocirrus which has a
strongly curled proximal trunk.
D. elongatus
Dordrechtites
DORDRECHTITALES
r/TRELITZIA 15 (2003)
65
Dordrechtites cetiparvus J.M.And. & H.M.And., sp. nov.
Holotype
Specimen : BP/2/1129; pi. 4(1, 6).
Assemblage (TC): Umk 1 1 1 Die 2spp; Umkomaas Valley.
Preservation : single dehisced scale, without counterpart; compression in
thinly laminated, carbonaceous (good cuticle) moderately baked, dark
grey shale with good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens'. 10 indivs (all isolated scales),
tfs 1-6 adjacent, pi. 4 (1-8).
Sister palaeodemes— nil.
Specific diagnosis
A Dordrechtites species bearing small scales with robust, very unequal
arms and sinuous, strongly reflexed fertile trunk.
Specific characters
Ovuliferous scale: small, compact; sterile arms robust, unequal, distal arm
markedly shorter (ca 5 mm long); fertile trunk relatively robust,
strongly curved proximally, sinuous towards apex.
Etymology
cetiparvus— cetus (Lat.), whale, after the locality Umkomaas (which means
cow or whale in the local Zulu language); parvus (Lat.), small, with
reference to the short arms of the scale.
Comments & comparison
The small size and curved arms of this species (known only from Umk
111) are somewhat reminiscent of the immature attached scales from Lut
311. However, as the scales are all found detached and isolated they are
regarded as mature and, therefore, as a distinct species.
Dordrechtites niazocirrus J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/4724a.b; pi. 4(15. 16).
Assemblage (TC): Maz 211 Hei/Dic; Mazenod.
Preservation: intact fascicle of four scales, part and counterpart; compres-
sion in thinly laminated, carbonaceous (poor cuticle) medium grey
shale with moderate cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: >40 indivs (2 clusters of 3-10 scales, >38 isolated scales),
tfs 7-11 adjacent, pi. 4 (9-16).
Sister palaeodemes— nil.
Specific diagnosis
A Dordrechtites species bearing relatively large scales with clearly
unequal arms and full, rounded, strongly reflexed fertile trunk.
Specific characters
Ovuliferous scales: relatively large; sterile arms gracile, unequal, distal
arm markedly shorter (ca 10 mm); fertile trunk robust, very strongly
arched proximally.
Etymology
mazocirrus—mazo (Lat.), after the locality Mazenod; cirrus (Lat.), curl,
with reference to the curled trunk of the scale.
Comments & comparison
The consistent nature of the rounded, strongly reflexed ovuliferous
trunk of the scales (known only from Maz 211), makes this species quite
unique.
D. niazocirrus
PRE/F/4696
pi. 4(10)
DORDRECHTITALES
Dordrechtites
66
d^TRELITZIA 15 (2003)
PRE/F/12760b
Aasvoelberg
(Aas 411 Dic/Sph)
PRE/F/20882
PRE/F/21747a
PRE/F/21762a
pl.l
Dordrechtites elongatus
DORDRECHTITALES
TRELITZIA 15 (2003)
Aasvoelberg
(Aas 311 Hei elo)
PRE/F/22515
PRE/F/22509
PRE/F/22512
1
%
X40
PRE/F/20864
PRE/F/18967
j f'M&A
PRE/F/22510jx2
10
DORDRECHTITALES
Dordrechtites elongatus
d?TRELITZIA 15 (2003)
Birds River
(Bir 111 Sph 2spp)
Lutherskop
(Lut 3 1 1 Hei elo)
Dordrechtites elongatus
DORDRECHTITALES
69
TRELITZIA 15 (2003)
PRE/F/8715
Umkomaas Valley
(Umk 1 1 1 Die 2spp)
BP/2/1133
PRE/F/4703
PRE/F/4696
PRE/F/4725
PRE/F/11750
BP/2/ 1129
BP/2/1133
BP/2/ 1128
BP/2/1129
Holotype
D. mazocirrus
Mazenod
(Maz 211 Hei/Dic)
PRE/F/4703
PRE/F/6782
PRE/F/8715
D. cetiparvus
pi. 4
DORDRECHTITALES
Dordrechtites
70
dA'RELITZIA 15 (2003)
PINOPSIDA S.V.Meyen 1984
VOLTZIALES J.M.And & H.M.And., ord. nov.
INCERTAE SEDIS family
Frediailthus J.M.And. & H.M.And., gen. nov.
Type species
Fredianthus maysiformis J.M.And. & H.M.And., sp. nov.
Generic diagnosis
A voltzialean male cone of very large size ( ca 220 mm long) with
strongly sigmoidal microsporophylls bearing a single row of ca 5 abaxial
clusters (ca 9 sacs each) of microsporangia.
Generic characters
Strobiius: a compact cone, lanceolate, of very large size (ca 220 X 53 mm
as reconstructed); axis stout (ca 11 mm diam.), moderately flexed;
microsporophylls helically attached, in ca 60 gyres of ca 25-30 units.
Microsporophylh. a simple scale, proportionately long, strongly sigmoidal,
at 90° from axis; distal lamina leafy, narrowly ovate, margin deeply
dentate with ca 12 lobes, without heel; stalk terete, narrow (ca 1 mm
diam.); microsporangia numerous (ca 45), abaxial, in a single median
row of ca 5 clusters, each with ca 9 radiating pollen sacs borne on a
short pedicel.
Microsporangium: elliptical (2x1 mm), retuse with clear, central, longi-
tudinal line (dehiscence slit?).
Pollen: unknown.
Eponymy
Fredianthus— in honour of Fred Terblanche, farmer at Goedehoop who first
showed us the type locality and has, more than once, aided excavation
with his tractor and ripper.
Global Range: Gondwana, Tr. (CRN).
First & last: the single Molteno species described here.
Gondwana Triassic occurrence
SAf— Karoo Basin, 1 TC (2 indivs).
Molteno occurrence
Frequency (F): 1 TC (of 100 sampled in the Molteno).
Diversity (D): 1 species.
Abundance (A): 2 individuals total.
Aas 411: 2 indivs in 512 man-hrs cleaving (1 per 25 man-days) extremely rare
Affiliated organs
Female: unknown.
Foliage: unknown.
With 13 ovulate genera recorded (Tab. 11, p. 15), Aasvoelberg (Aas
41 1 ) is the richest fruit-yielding TC in the Molteno. Yet there occurs at the
site no obvious ovulate coniferopsid cone that might affiliate with
Fredianthus. Dordrechtites, without a known affiliate, must be considered
a remote possibility.
A similar problem arises with regard to a foliage affiliate. Three conif-
eropsid genera —Heidiphyllum (1%), Rissikia (25 individuals) and
Pagiophyllum (two small fragments)— occur in the assemblage, none of
which is a likely candidate. Heidiphyllum and Rissikia are preoccupied,
while Pagiophyllum (known nowhere else in the Molteno) is considered to
belong to the order Pinales and not the more primitive Voltziales.
Classification & comparison
Suprageneric classification (family Incertae/Voltziales)
Fredianthus is placed in the Voltziales— transitional between the
Cordaitanth.ales and Pinales— though it is quite unlike any of the Laurasian
male cones included in the order (pp. 57, 59). It is perhaps most readily
derived from a form such as the Siberian Permian genus Cladostrobus
(Rufloriaceae, Cordaitanthales, tf. 25, p. 59). In view of the lack of any
established affiliation with ovulate cones or foliage, Fredianthus is not
included in any formal family. However, considering its unique size and
arrangement of microsporangia, it probably represents a new group at this
rank.
Intergeneric comparison (Molteno genera)
Odyssianthus and Lutanthus, sister genera of Fredianthus in the
Molteno, are judged sufficiently distinct— given what is currently known
of male conifer cones — to represent separate families within the Voltziales
(Tab. 30, p. 54). See Tab. 31 (p. 57) for a possible resolution of the rela-
tionships between the male pinopsid cones from the later Carboniferous to
Triassic.
Reconstruction
Though two specimens of this remarkable cone exist in the collection,
the reconstruction is based on the holotype alone— the second individual
being a small, poorly preserved fragment. The apex and base of the cone,
being unknown, are conjecture. The number of scales per gyre, the distal
lamina and the arrangement and number of microsporangia are a best attempt
at interpretation, the security of which will clearly increase with more and
better preserved material. In view of the extreme rarity of the taxon, however,
the gathering of additional specimens is not too likely in the near future.
Fredianthus
VOLTZIALES
c/tRELITZIA 15 (2003)
71
Fredianthus maysiformis j.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/1 2753a, b; pi. 5(1-4), pi. 6(1-7).
Assemblage (TC): Aas411 Dic/Sph; Aasvoelberg.
Preservation : part and counterpart of a fairly complete cone (175 X 55
mm); apex and proximal end with stalk missing; preserved primarily
in longitudinal section with outer face of scales seen only in variable
side view along cone margin; microsporangia in situ and undehisced
throughout; impression in thinly laminated, strongly baked, yellowish
grey shale with very good cleavage.
Reference palaeodeme
Assemblage (TC); as for holotype.
Specimens: 2 individuals; including the holotype and an additional small
cone fragment.
Sister palaeodemes — nil.
Specific diagnosis— as for genus.
Specific characters— as for genus.
Etymology
maysiformis— with reference to the superficial similarity to a maize (com,
mealie) cob.
Classification & comparison
Fredianthus maysiformis , at ca 200 mm length, appears to be the
largest Pinopsida male cone— fossil or extant— yet recorded. In the extant
Coniferales, the male cones are generally small (30 mm long or shorter)
and the sporangia few in number. Throughout the Pinaceae there appear
only two sporangia, and in most other extant families from two to seven
sporangia. The largest male cone ( Araucaria bidwilli) reaches a length of
120 mm and the highest number of sporangia per sporophyll (in Agathis ) is
13-15 (Foster & Gifford 1974).
In contrast, the Cycadales bear numerous sporangia (30 in Zamia to
>1 000 in Cycas ) and have characteristically large male cones (>200 mm).
A very large cone (200 mm long), presumed to be female, is recorded from
the Burgersdorp Fm. as Sewardistrobus (And. & And. 1985).
S
VOLTZIALES
Fredianthus
72
c/tRELITZIA 15 (2003)
all PRE/F/12753a Holotype
Aasvoelberg
(Aas 411 Dic/Sph)
VOLTZIALES
Fredianthus maysiformis
73
TRELITZIA 15 (2003)
all PRE/F/12753a Holotype
VOLTZIALES
Fredianthus maysiformis
74
d?TRELITZIA 15 (2003)
PINOPSIDA S.V.Meyen 1984
VOLTZIALES J.M.And. & H.M.And., ord. nov.
INCERTAE SEDIS J.M.And. & H.M.And., fam. nov.
Lutanthus J.M.And. & H.M.And., gen. nov.
Type species
Lutanthus hemidiscus J.M.And. & H.M.And., sp. nov.
Generic diagnosis
A voltzialean male cone of small size (ca 30-40 mm long) with
straight to moderately upcurving microsporophylls bearing few microspo-
rangia (6-8) in diverse modes of attachment.
Generic characters
Strobilus: a compact cone, oblong-elliptic, apex rounded, base truncate, of
relatively small size (ca 30—40 mm long); axis stout to very stout,
strongly flexed to erect, without free base (cone sessile); microsporo-
phylls helically attached, number of gyres (13-24) and units per gyre
(10-20) very variable.
Microsporophyll. a simple scale, proportionately short, curving upward
distally, at 45° to 90° from axis; distal lamina leafy, ribbed to lobed to
hemispherical and entire, without heel; stalk narrow to broadly taper-
ing, variously winged; microsporangia few in number (6-8), adaxial to
abaxial, in 1 or 2 rows, clustered or single, sessile.
Microsporangium: variously obovate to elliptical or rhomboidal (ca 1 mm
long), generally with a clear longitudinal (?)dehiscence slit.
Pollen: unknown.
Etymology
Lutanthus— after the type locality, Lutherskop.
Global Range: 3 spp., Gondwana, Tr. (CRN).
First & last: the three Molteno species described here.
Gondwana Triassic occurrence
SAf— Karoo Basin, 3 TCs (5 indivs).
Molteno occurrence
Frequency (F): 3 TCs (of 100 sampled in the Molteno).
Diversity (D): 3 species.
Abundance (A): 5 individuals total, very rare to vanishingly rare.
Gre 111 Equ sp: 2 indivs in 25 man-hrs cleaving (1 per 1 man-day) very rare
Lut 311 Hei elo: 2 ” ” 50 ” ” (1 ” 2'L ” ) very rare
Aas 411 Dic/Sph: 1 ” ”512 ” ” (1 ” 51 ” ) vanish, rare
This pattern of scarcity recorded for Lutanthus is consistent with that
noted for the other Molteno voltzialean male cones, Fredianthus and
Odyssianthus.
Affiliated organs
Female cone: unknown.
Foliage: unknown.
Though we recognise in the Molteno only a single genus each of
voltzialean foliage ( Heidiphyllum ) and female cone (Telemachus), there
appear to occur, in the very scarce material at hand, at least three genera of
male cone. For neither Lutanthus or Fredianthus are there further foliage or
female taxa (in the Pinopsida) in the collections for suggesting possible
affiliations.
Classification & comparison
Suprageneric classification (Incertae Sedis/Voltziales)
Lutanthus, along with its two Molteno sister genera, Fredianthus and
Odyssianthus, is included, with considerable reservation, in the order
Voltziales (see discussion on pp. 70, 88). In the absence of affiliated ovu-
late cones or foliage, the genus is placed as family incertae sedis. It would
be quite feasible to derive Lutanthus from the Late Permian glossopterid
microsporangiate genus Lidgettonia as depicted in Tab. 3 1 (p. 57) and
noted on p. 56— offering a wholly different polyphyletic pinopsid phy-
togeny.
Intergeneric comparison (Molteno genera)
In what measure do the three Molteno voltzialean male cones,
Lutanthus, Fredianthus and Odyssianthus, differ? They are represented by
a total of only nine individuals and could conceivably, at one end of the
spectrum of possibilities, be placed in a single genus. The fact that the
Molteno voltzialean foliage is confined almost exclusively to the highly
dominant genus Heidiphyllum tends to support this option. We favour the
three-genera alternative, however, for the reasons best clarified in the text,
tables and comparative figures on pinopsid phytogeny and morphology, pp.
56-59. Indeed, we find it most plausible within the context of the great
Triassic radiation that three distinct families, possibly even orders, might
be represented.
Lutanthus
VOLTZIALES
C^TRELITZIA 15 (2003)
75
assemblages
(taphocoenoses)
Ct, Fredianthus
Q, Lutanthus
°» Odyssianthus
Q, Rissikianthus
O. Helvetianthus
1
Gre 111 Equ sp
- 2 - - -
2
Boe 112 Die cor
. . . i -
3
Tel 111 Hei elo
. - 2 - -
4
Lut311 Hei elo
- 2 - - -
5
Pen 321 Dic/Ris
- - - 30 -
6
Kap 111
- - - 25 -
7
Umk 111 Die 2spp
- - - 8 -
8
Lit 111 Dic/Hei
- - - - 6
9
Aas411 Dic/Sph
21 - 15 -
Total TCs
13 15 1
Total indivs
2 5 2 79 6
Tab. 34. Pinopsid pollen cone genera,
Molteno occurrence
Matrix: curated individuals in collection
Frequency/abundance: aside from Rissikianthus,
it is clear that pinopsid pollen cones are
rarely encountered
Adaptive radiation
The rather disparate group of three species from three different
Molteno localities is brought together into the single genus Lutanthus with
obvious uncertainty since the material is so scarce and none of it is com-
pletely or explicitly enough preserved to render all diagnostic features
unambiguous. The differences between the three species are so marked that
we might well be dealing with three distinct genera; but if so, where are
their foliage and female affiliates? The most diagnostic features differenti-
ating the species are seen in the various elements of the scales— distal lam-
ina, stalk and pollen sacs.
Reconstructions
The R4 reconstructions of the three Lutanthus species portrayed here
are variously secure; based as they are on so few (five in total) individuals.
It is most particularly in the arrangement and attachment of the microspo-
rangia that the uncertainties lie. Settling decisively on the critical question
of adaxial or abaxial emplacement has proved notoriously tricky for all
three species. The options as illustrated and described seem most likely, but
are not certain. To summarise: the microsporangia in the three species
appear to range from being attached in two latero-abaxial rows (in L. orna-
tus), to two latero-adaxial rows ( L . hemidiscus ), to a single adaxial row (L.
robustus). If one allows for the lateral wings of the stalks being suppressed
and a small degree of adaxial migration of the two rows of microsporangia
in the first two species, then a similar morphological condition in the three
species can be visualised. The trio of species being related at generic level
is conceivable (but see also text adjacent).
From the material at hand (pis 78-81), which lacks any evidence of
free axes, we take all three species to be sessile.
The three Molteno species are based on the following TCs/reference
palaeodemes. Each derives from a very distinct habitat and from a differ-
ent member within the stratigraphic sequence.
L. hemidiscus— hut 311 Hei elo (Lutherskop); 2 indivs
Heidiphyllum thicket; Cycle 3 (Mayaputi Member)
L. ornatus— Gre 111 Equ sp. (Greenvale); 2 indivs
Equisetum marsh; Cycle 4/5 (Qiba Member)
L. robustus — Aas411 Dic/Sph (Aasvoelberg); 1 indiv
Sphenobaiera closed woodland; Cycle 1 (Bamboesberg Member)
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
SAm SAf Ind Ant Aus
Ch NA SA Pa Lu | Za 1 Li | Ka WH I PI A* NZ Ca SA Ga j Bo CM
VOLTZIALES
Lutanthus
76
d^TRELITZIA 15 (2003)
Lutanthus hemidiscus J.M.And. & H.M.And., sp. nov.
Holotype
Specimen : PRE/F/1 1537a,b; pi. 7(1-5).
Assemblage (TC): Lut 311 Hei elo; Lutherskop.
Preservation : fairly complete cone, part and counterpart, longitudinal
outer view with several scales and numerous microsporangia clearly
evident; impression in thickly laminated, medium grey shale with
moderate cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens : 2 indivs (tfs 1, 2 opposite).
Sister palaeodemes— nil.
Specific diagnosis
A Lutanthus cone with strongly flexed axis, large hemispherical scales,
and microsporangia with distinctive (?)dehiscence line.
Specific characters
Strobilus: axis moderately stout, strongly flexed to base; microsporophylls
in ca 13 gyres of ca 10-12 units.
Microsporophyll: distal lamina large, hemispherical, margin entire; stalk
broad, spreading distally, widely winged; microsporangia adaxial, dor-
solaterally attached to midrib, single, sessile, in two irregular rows of
3 or 4 pollen sacs.
Microsporangium: ca 1.1 mm long, variously obovate to elliptical, occa-
sionally kidney-shaped; with distinct (?)dehiscence line running full
length.
Etymology
hemidiscus (Lat.)— half disc, with reference to the shape of the scale.
Comment & comparison
With two relatively complete individuals from Lut 311, the holotype
seen in rather disarticulated outer view, the sister specimen in section, L.
hemidiscus is the best known of the three Lutanthus species and is the obvi-
ous choice as type species for the genus. Its conspicuous, hemispherical
scale laminae with entire margins are particularly characteristic.
Lutanthus ornatus J.M.And. & H.M.And., sp. nov.
Holotype
Specimen : PRE/F/16534; pi. 9(1-6).
Assemblage (TC): Gre 111 Equ sp.; Greenvale.
Preservation : nearly complete cone (proximal end missing), without coun-
terpart, longitudinal outer view and part section, with a couple of
scales and numerous microsporangia clearly evident; impression in
thickly laminated, medium grey shale with good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 2 indivs (tfs 9, 10 opposite).
Sister palaeodemes — nil.
Specific diagnosis
A Lutanthus cone with straight axis, distinctively lobed scales, and
ornamented microsporangia with clear ‘micropyle’.
Specific characters
Stobilus: axis moderately stout, expanding strongly to base, not flexed;
microsporophylls in ca 17 gyres of ca 15 units.
Microsporophyll: distal lamina fairly large, distinctively multilobed; stalk
linear, narrowly winged; microsporangia abaxial, ventrolaterally
attached to midrib, single to semiclustered, sessile, in 2 rows of ca 4
pollen sacs.
Microsporangium: ca 1 .1 mm long, obovate, oval to circular in end view;
with distinct linear-obovate (?)dehiscence line running most of length;
and a distinctive, linear cellular ornamentation radiating out from api-
cal ‘micropyle’.
Etymology
ornatus (Lat.)— with reference to the ornamented microsporangia.
Comment & comparison
L. ornatus , with its clearly ornamented microsporangia and well-lobed
scales, is as distinctive a species as L. hemidiscus. The lobed scale laminae
are, however, confusingly camouflaged amongst the mass of pollen sacs in
the two available specimens. It is the difference in ornamentation between
lamina and sacs, as seen under the microscope, that sets these two elements
of the cone apart.
Lutanthus
VOLTZIALES
^YtrELITZIA 15 (2003)
77
Lutanthus robustus J.M.And. & H.M.And., sp. nov.
Holotype
Specimen'. PRE/F/20876a,b; pi. 10(1-5).
Assemblage (TC): Aas 41 1 Dic/Sph; Aasvoelberg.
Preservation: Fairly complete cone (proximal end missing), part and
counterpart, longitudinal section showing numerous scales and a scat-
ter of in situ microsporangia; impression, imperfectly preserved;
impression in thinly laminated, strongly baked, yellowish grey shale
with very good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 1 indiv. (tf. 1 opposite).
Sister palaeodemes— nil.
Specific diagnosis
A Lutanthus cone with gently flexed robust axis, small ribbed scales,
and rhomboidal microsporangia without distinctive features.
Specific characters
Strobilus: axis markedly robust, very gently flexed to base; microsporo-
phylls in ca 24 gyres of ca 20 units.
Microsporophyll: distal lamina relatively small, kidney-shaped, multi-
ribbed, margin dentate-crenate; stalk linear, moderately winged;
microsporangia adaxial, dorsally attached to midrib, in 2 sessile radi-
ating clusters of 4 or 5 pollen sacs.
Microsporangium: ca 0.8 mm long, irregularly rhomboidal; with no evi-
dent mode of dehiscence or ornamentation.
Etymology
robustus (Lat.)— with reference to the stout axis.
Comment & comparison
L. robustus, based on only one individual from Aas 411, is the least
clearly preserved of the three species grouped here under Lutanthus, and
is, perhaps, the most different. Its particularly stout axis, small microsporo-
phyll laminae, and adaxial clusters of rhomboidal pollen sacs, set it apart.
In bearing a single row of microsporangial clusters— though these are
adaxial, not abaxial, and with very different sacs— L. robustus is reminis-
cent of Fredianthus (pp. 70. 71). The multiribbed scale laminae also show
similarities. On these counts, the two taxa could conceivably be congener-
ic, yet the differences seem more profound than the similarities. The
species is placed provisionally under Lutanthus, but with obvious reserva-
tion.
L. robustus
Aas 4 1 1
PRE/F/20876a,b
pi 10(1-5)
Holotype
all x2'/2
unless indicated
L. hemidiscus
end view
12
w
- xlO
pi. 7(2,4, 5)
ly xlO
Lut 311
PRE/F/1 1 537a,b microsporangia all R2
L. ornatus
ri
Tit
W%
Gre 111
PRE/F/16534 microsporangia all R2
L. robustus
O'.
Q/\
Oft
''l
„ ogr
pi. 10(2)
tjjy
pi. 10(3-5)
Aas 411
PRE/F/20876a,b microsporangia all R2
VOLTZIALES
Lutanthus
pi. 7
Lutanthus hemidiscus
VOLTZIALES
r''/ TRELITZIA 15 (2003)
pi. 8
VOLTZIALES
Lutanthus hemidiscus
80
<S?TRELITZIA 15 (2003)
Lutanthus ornatus
VOLTZIALES
all PRE/F/16534 Holotype
Greenvale
(Gre 1 1 1 Equ sp)
g]
TRELITZIA 15 (2003)
VOLTZIALES
pi. 10
Lutanthus robustus
82
TRELITZIA 15 (2003)
PINOPSIDA S.V.Meyen 1984
VOLTZIALES J.M.And. & H.M.And., ord. nov.
VOLTZIACEAE Florin 1951
Telemachus H.M.And. 1978
Type species
Telemachus elongatus H.M.And. 1978.
Generic diagnosis
A voltzialean female cone with megasporophylls bearing a pair of
adaxial ovules on a 5-lobed ovuliferous scale.
Generic characters
Attachment : terminal, sessile, erect, occurring singly on woody stem.
Strobilus : simple, compact cone, oval, moderate size (25-75 mm long);
axis stout, erect, without free base; megasporophylls helically
attached.
Megasporophyll: bract/scale complex extends from axis at right angles;
sterile bract fused to ovuliferous scale for half its length, distal end
free, leafy (7-45 mm long), often curving strongly to base, linear to
lanceolate, proximal margin finely dentate, tapering to acute tip;
ovuliferous scale bearing 2 adaxial ovules, distally strongly upturned
with 5 distinct acute lobes.
Ovule/seed: naked, oblong to oval (ca 4x2 mm).
Eponymy
Telemachus— the son of Odysseus and Penelope in Greek mythology; after
the holotype locality of the type species.
Global range: 7 spp. Gondwana, Tr. (LAD-CRN).
First : Telemachus lignosus (Retallack 1981b); Long Gully Fm., Benmore
Dam region. New Zealand.
Last: Telemachus elongatus (Anderson 1978); Molteno Fm.
Gondwana Triassic occurrence
SAm— N. Argentina, 2 Iocs (4 indivs).
SAf —Karoo Basin, 18 TCs (>300 indivs).
Ant -Trans-Antarctic Mts, 2 Iocs (5 indivs).
Aus —New Zealand, 1 loc. (4 indivs).
Molteno occurrence
Frequency (F): 18 TCs (of 100 sampled in the Molteno).
Diversity (D): 6 species.
Abundance (A): 311 individuals total, abundant (at Gre 121 Hei elo) to
very rare (at Pen 321 Dic/Ris).
Qac 1 1 1 Hei/Dic: 10 indivs in 4 man-hrs cleaving (25 per 1 man-day) rare
Aas 611 Hei elo:
6 ”
” 3
” (20 ”
Kle 1 1 1 Hei elo:
25 ”
” 15
(20 ”
Pen 411 Hei elo:
>50 ”
” 70
” (10 ”
Kan 112 Hei elo:
10 ”
” 15
” (10 ”
Tel 111 Hei elo:
40 ”
” 90
” ( 5 ”
We include data above only for the reference TCs of the six Molteno
species described. In that the figures are based exclusively on curated spec-
imens, the relative abundance of Telemachus will be generally under-
played. For details of the frequency and abundance of the three affiliated
genera, Telemachus (female), Odyssianthus (male) and Heidiphyllum
(foliage), through the Molteno. see Tab. 36 (p. 84). For analysis of the
mother plant’s preferred habitat, and of a typical TC (Aas 311 Hei elo), in
which it occurs as a monodominant, see And. & And. (in prep).
Affiliated organs
Male strobilus: Odyssianthus— Grade 4 (Kin. reinf., Mut. occ.).
Foliage: Heidiphyllum— Grade 4 (Kin. reinf., Mut. occ.).
Evidence for affiliation between the three genera Telemachus , Odyssian-
thus and Heidiphyllum is convincing both regarding kindred reinforcement
and mutual occurrence (see further on pp. 84, 88).
Classification & comparison (see also Odyssianthus)
Suprageneric classification (Voltziaceae/Voltziales)
Telemachus and Heidiphyllum both fall readily within the morphologi-
cal ambit of the Voltziaceae as defined in And. & And. ( 1 989, pp. 422. 423).
Podozamites, a common and widespread Laurasian Late Triassic to Early
Jurassic genus, superficially very similar to Heidiphyllum, is known in
organic connection with the genera Borysthenia and Cycadocarpidium and
in close affiliation with Swedenborgia. These three northern genera are typi-
cal female strobili of the Voltziaceae.
Intergeneric comparison (Gondwana Triassic)
Telemachus stands as a very clearly defined genus and the only female-
cone representative of the Voltziales in the Gondwana Triassic.
Reconstructions
The pen sketches (tfs 1-3 above) depicting the type species T. elonga-
tus (from Tel 111), with its long, narrowly lanceolate reflexed bracts are
Grade 5 reconstructions (R5). They are based on the full reference
palaeodeme (pi. 11, figs 1-11) for the species, along with supporting evi-
dence from the sister palaeodeme Aas 111 (pi. 12, figs 10-12).
Bract/ scale complex: Two grades of line drawing have been prepared for
each species (pp. 85-87): a number of R2 sketches showing the specimens
as preserved; and an R4 sketch depicting a complete bract/scale complex
representing the norm of the reference palaeodeme.
Molteno occurrence (elaborated)— extracted from And. & And. (in prep.)
Mother-plant ( Heidiphyllum/Telemachus/Odyssianthus)
The Heidiphyllum elongation mother plant is seen as an erect woody
shrub up to 3 m, forming near-monospecific stands in waterlogged habitats.
Foliage ( Heidiphyllum )
Heidiphyllum , occurring in 62 TCs, is the second most frequent and
abundant genus in the Molteno. In the Heidiphyllum-thicket TCs it often
overwhelmingly dominates the assemblage to the extent where all other
taxa together amount to less than 1 or 2% of the total. Where it occurs com-
monly in TCs of other vegetation types, we visualise the genus as deriving
from similar monodominant thickets but of varying extent and varying dis-
tance further afield.
Female cone ( Telemachus )
In strong contrast to Odyssianthus, the female, Telemachus, either
intact or as partial cones or detached scales, is remarkably common. The
pattern is clear that this strong presence is almost exclusively associated
with the Heidiphyllum- thicket TCs. Here it occurs in over half the total
number of assemblages.
Male cone (Odyssianthus)
The male remains exceptionally rare, with only two cones having been
found in a single TC, Telemachus Spruit (Tel 111). Remarkably, these
cones are intact, more or less complete, and with in situ microsporangia.
Telemachus
VOLTZIALES
^TRELITZIA 15 (2003)
83
Gondwana Triassic occurrence (elaborated)
Although Heidiphyllum is one of the most prominent genera (frequen-
cy and abundance) throughout the Gondwana Triassic— often a monodom-
inant overwhelming assemblages— its female and male cones (outside of
the Molteno) remain extremely rare and absent respectively. A total of only
12 Telemachus individuals, all intact strobili, from South America,
Antarctica and New Zealand are recorded in the literature (Tab. 35).
Remarkably, the cone has not yet been recorded from Australia.
South America (Frenguelli 1942; Spalletti etal. 1991)
Frenguelli (1942) described a new genus, Pterorrachis, with two
species, P. ambigua and P. problematica, based on two intact cones from a
locality near Ischigualasto, San Juan Province, N. Argentina. The photos of
the specimens are poor and the reconstruction sketches unconvincing. The
material is tentatively identified here as Telemachus— an identification sup-
ported by the associated occurrence of Heidiphyllum ( Phoenicopsis in
Frenguelli 1942). A distinctive, complete Telemachus cone (?species) was
found by HMA (22-09-1999) on a field trip (VII International Symposium
on Mesozoic Terrestrial Ecosystems) to Ischigualasto— El Gusano locality,
Los Rastros Fm. It was given to Alfredo Monetta for their collection at San
Juan.
A fourth (possible) cone was recorded and illustrated by Spalletti et al.
(1991) from Arroyo Lapa, Lapa Fm., Neuquen Basin, central Argentina,
and identified as Telemachus elongatus. The specimen is intact, but very
unclear and not identifiable to species.
Antarctica (Yao et al. 1993; Axsmith et al. 1998a)
A total of five intact Telemachus strobili (compressions) have thus far
been recorded from two widely separated localities, Mount Falla and Allan
Hills, in the Triassic strata of the Transantarctic Mountains.
The Mount Falla locality (Yao et al. 1993) occurs in the Queen
Alexandra Range (Buckley Island Quadrangle) within the Lower Falla Fm.
of the latest Ladinian. Though intact and clearly preserved, the two Mount
Falla specimens do not show the diagnostic bract/scale features and are
thus included in the hypodigm table under T. spp. indet. Yao et al. note that
numerous specimens of Heidiphyllum are ‘found together with Telemachus
in the Falla Formation’.
The three Alan Hills specimens (Axsmith et al. 1998b) from the Lashly
Fm. are likewise intact and clearly preserved, yet do not adequately show
the diagnostic bract/scale features. In this case, since one of the cones does
at least show elongate bract scales, we include the three-specimen
palaeodeme tentatively as T. elongatus.
New Zealand (Retallack 1981b)
Telemachus lignosus, described by Retallack (1981b) on the basis of
four intact cones, derives from cuttings along the Backyards to Otematata
River road (S 1 17/F754), Long Gully, Long Gully Fm., Upper Corbies
Creek Group, Kaihikuan Stage, Ladinian. The specimens are not clearly
preserved, but from the pen sketches and reconstruction of Retallack, the
bract/scale complex, with short bracts and scales lacking obvious outer
angles (elbows), appears sufficiently different from any of our Molteno
taxa to warrant recognition as a distinct species. The low-diversity TC
yielding 71 lignosus includes Heidiphyllum , but Retallack does not record
relative abundance data.
Tab. 35.
TELEMACHUS HYPODIGM, Gondwana Triassic occurrence
Species
Intact-
ness
Molteno
Other
T. elongatus
T. grandis
T. serribractus
T. brachybract.
T. dubibractus
T. acutisquamus
T. lignosus
T. spp. indet
Intact strobili
Fragmentary ”
Isolated scales
AUTHOR
SUBREGION (“square)
FORMATION
LOCALITY
NAME (original)
Indivs ILLUS.
SOUTH AMERICA
1942 Frenguelli
1991 Spalletti et al.
Ischigualasto NA1
Paso Flores SA1
? ?
? ?
24 Lapa Fm.
nr. Ischigualasto
Arroyo Lapa
Pterorrachis ambigua
" problematica
Telemachus elongatus
1 pi 1(1), tf 1
1 pi 1(2,3), tf 2
1 pi 3(a)
- 1
- 1
- 1
1 - -
1 - -
1 - -
ANTARCTICA
1 993 Yao et al.
1 998a Axsmith et al.
Beardmore Glacier TA4
Allan Nunataks TA1
22 L. Falla Fm.
22 Lashly Fm.
Mount Falla
Allan Hills
Telemachus elongatus
2 pi 1(1-5)
3 pi on p710(18-20)
3
- 2
2 - -
3 - -
NEW ZEALAND
1981b Retallack
Benmore Dam NZ4
21 Long Gully
Long Gully
Telemachus lignosus
4 pi 11(J-L), tf 5(A-E)
4 -
4 - -
SOUTH AFRICA
1978-1999: And. & Ar
d. Molteno literature not inc
luded in this tabl
e
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
VOLTZIALES
Telemachus
84
dA’RELITZIA 15 (2003)
Evidence for affiliations
Foliage : Heidiphyllum— Grade 4 reliability
As in the case of four further Molteno female strobili,
Umkomasia, Peltaspermum, Rissikistrobus and Fraxinop-
sis , the foliage affiliate of Telemachus is almost certainly —
short of organic attachment— established. The foliage
genus Heidiphyllum, abundant and widespread throughout
Gondwana during the Middle and Upper Triassic, occurs in
close mutual occurrence with Telemachus in three, perhaps
four, continents in which the latter is now known.
South Africa (Tab. 36)— Telemachus is known from 18
of the 100 Molteno TCs sampled. Heidiphyllum occurs in
all of these TCs, in 14 cases as a co-dominant to dominant
element of the assemblage.
South America (Frenguelli 1942 )—Phoenicopsis (i.e.
Heidiphyllum ) reportedly occurs on the reverse of both fos-
siliferous slabs bearing possible Telemachus.
New Zealand (Retallack 1981b)— The single low-
diversity TC yielding the four recorded specimens of
Telemachus includes the genus Heidiphyllum. Retallack
does not record the relative abundance of the foliage taxa
occurring at the site.
Antarctica (Yao et at. 1993)— Telemachus is found in
association with numerous specimens of Heidiphyllum in
the Falla Formation.
Cuticular correspondence
The only available cuticular data on Telemachus are
those of Yao et al. (1993), who illustrate and describe a
number of fragments from Antarctic material, which they
consider well preserved but in our scheme we rate fair
(Grade 3).
In our revision of Molteno gymnosperm foliage (And.
& And. 1989. pp. 54, 55), Heidiphyllum cuticle fared poor-
ly—with only three of the 23 leaves sampled for maceration
from Lit 111 and Umk 111 yielding fragments of fair
(Grade 3) preservation. Most revealed poor (Grade 2) to
very poor (Grade 1) preservation. The best cuticular results
obtained from Umk 111 and Lit 111 complement one anoth-
er, the former based on a macerated specimen, the latter on
a relatively transparent cellulose-acetate peel.
On the basis of the incomplete results at hand, the com-
parison between the cuticle of Heidiphyllum and Tele-
machus is suggestive but inconclusive. The oblong to pen-
tagonal or hexagonal cells with straight to gently curved
walls are similar, as are the five or six noncutinised, anomo-
cytic subsidiary cells surrounding the stomata. Strongly
cutinised guard-cells appear to be a feature of both taxa. but
are not always preserved. The bold papillae of the lower
cuticle and the stomatal lappets in Heidiphyllum have not
been observed in Telemachus.
Male strobilus: Odyssianthus— Grade 3 reliability
The male counterpart of Telemachus is extremely infre-
quent and rare in the Molteno. being known from only two
individuals from 1 TC (Tel 111). Heidiphyllum is the over-
whelming dominant (89%) in this assemblage.
Kindred reinforcement
Telemachus and Heidiphyllum both fall readily within
the morphological concept of the family Voltziaceae as
delimited and summarised in And. & And. (1989, pp. 422,
423). A generous range of female cone genera are included
in the family. Podozamites, a common and widespread
Laurasian U. Triassic to L. Jurassic foliage genus, superfi-
cially very similar to Heidiphyllum, is known in organic
connection with the genera Borysthenia and Cycadocarpi-
dium and in close affiliation with Swedenborgia. The latter
three genera are typical female strobili of the Voltziaceae.
Tab. 36. TelemachusIHeidiphyllum, Molteno occurrence
assemblage
(taphocoenosis)
Nav 111
Die odo
1
Cal 211
Hei elo
75
Bir 211
Sph 2spp
3
-
311
Hei/Sph
45
-!
111
Sph 2spp
10
Dor 111
Hei elo
85
-i
Gre 121
Heo elo
98 100
111
Sph pon
10
-1
Boe 111
Lep sto
7
-!
Dic/Hei
42
-!
112
Die cor
14
-i
Cyp 111
Die era
24
5
111
Hei elo
100
- 1
Kan 112
Hei elo
98
10
Ast spA
10
-!
Tel 111
Hei elo
89
40
Vin 111
Die odo
28
_l
Ela 111
Die odo
7
1
Kra 311
Die odo
5
-!
Lut 111
Hei/Dic
50
2!
511
Hei elo
80
1
411
Hei/Dic
50
12
311
Hei elo
99
-1
Tin 121
Sph 2spp
4
- 1
111
Sch sp.
10
-
131
Hei/Ast
50
1
Kon 223
Die odo
1
211
Hei elo
95
111
Die odo
7
-!
Sch sp
10
-1
Hei elo
84
2
Pen 321
Dic/Ris
1
1
211
Dic/Equ
2
-
421
Die odo
4
431
Dic/Equ
5
-
311
Hei elo
75
17
411
Hei elo
94
50
Kle 111
Equ sp
1
_ 1
Hei elo
90
25
Hei/Dic
49
- 1
Kap 111
Dic/Ris
25
5
Ela 112
Dic/Hei
30
Nuw 211
Die 2spp
1
_ 1
Win 111
Hei elo
79
-1
Qua 111
Die odo
20
Maz 111
Die era
5
“ 1
211
Hei/Dic
32
Hla 213
Die elo
1
1
Umk 111
Die 2spp
7
_ 1
Inj 211
Die dub
10
-!
San 111
Die era
5
- 1
Mng 111
Die 2spp
6
Qac 111
Hei/Dic
50
101
Mat 111
Die dub
4
12
Lit 111
Dic/Hei
23
-!
Aas 611
Hei elo
80
6
111
Hei elo
77
12
211
Hei elo
100
-]
311
Hei elo
99
411
Dic/sph
1
-!
511
Die elo
20
-!
Bam 111
Die dub
1
E
C </)
0)
£ o
! 9 ! cf
Species
3 t
re u>
y
40
yy
yy
y
10
io; 30
yyy
yyy
T~5
yy
yy
2b
2a
12
yyy \
Total TCs
Total indivs
62; 18; 1
% 311 2
1 21 2 1
10 1"37^ 67 M0
1 6
6 22
10 13
31 Tisi
Telemachus
VOLTZIALES
& TRELITZIA 15 (2003)
85
T. grandis
R4
\2
V
Kan 112
BP/2/3415 etal.
pi. 12(1-5)
T. serribractus
PRE/F/7693 et al. PRE/F/4328a
pi. 11(1-12) pi. 12(11)
T. acutisquamus
Telemachus
generic panorama showing
the 6 Molteno species
R4s based primarily on the
individuals indicated, but the full
set of illustrated specimens from
the reference palaeodeme is
considered.
T. dubibractus
Pen 411
PRE/F/17180 etal.
pi. 15(1-9)
Qac 111
BP/2/290 la etal.
pi 16(7-10)
pair of seeds
attached to scale
11
T. brachybr actus
y \ i \ dispersed seeds
vi
Pen 411
PRE/F/19632
pl. 15(8)
Pen 311
PRE/F/16941a <Xr&
pl 16(5)
Intactness of cones (Tab. 36)
Degree of cone fragmentation
A substantial proportion of Telemachus individuals are found as intact
or partially intact cones. While the proportions indicated in the table are
inevitably biased towards the more complete material, it is perhaps reason-
able to estimate that around 5% of Telemachus specimens overall are found
as more or less fully intact cones.
In situ seeds
Most Telemachus cones, partial cones or isolated scales found in the
Molteno Fm. have lost their seeds. In only seven of the 18 TCs yielding
Telemachus do any in situ seeds occur; in these seven TCs only a small pro-
portion (10% or less of individuals) still bear any seeds, and in those few
cones still bearing seeds, more often than not only a few seeds remain in
situ. A selection of cones from four TCs (mostly illustrated, pis 11-16)
showing different states of seed dispersal are noted below.
Kle 111 Hei elo (8 intact or fragmentary cones)— In this TC a particu-
larly interesting suite of clearly preserved cones occurs— BP/2/5889 is a
cone seen in cross-section with most seeds still in situ\ BP/2/5889 is a lon-
gitudinal section of cone with most seeds lost, while BP/2/5891 is a fine
cross-section of cone with all seeds lost (pl. 13).
Pen 411 Hei elo (40 intact or fragmentary cones)— This TC has yield-
ed the most comprehensive Telemachus palaeodeme as well as numerous
readily identified dispersed seeds. PRE/F/19632 (pl. 15(8)) is the clearest
specimen in the Molteno collection showing a pair of seeds attached to a
scale.
Kan 112 Hei elo (2 intact or fragmentary cones)— PRE/F/20080a,b is
the sole instance in the Molteno collection where the cone is found
attached— terminally on a short section (2 cm) of stout broken shoot (pl.
12(1-3)). A few seeds remain in situ in this specimen.
Lut 411 Hei/Dic (5 intact or fragmentary cones)— PRE/F/ 147 15 is a
partial cone with a particularly full complement of in situ seeds (not illus-
trated).
Dispersed seeds
Dispersed seeds are recorded at a further five TCs aside from the four
listed above. It should be noted that only those TCs yielding Telemachus
have been checked for scattered seeds and most of these bear the dis-
seminules in fair numbers.
Adaptive radiation (Molteno diversity)
Telemachus (female strobilus)
As in at least two other notable Molteno plant-genera. Peltaspermum/
Lepidopteris and FraxinopsislYabeiella, diversification in Telemachus /
Heidiphyllum is more evident (morphologically manifested) in the female
strobilus than in the male counterpart or the foliage. It is in the bract/scale
complexes, and in particular the shape, marginal dentition and size of the
bracts, that the signature of the six species is best portrayed (tfs 1-9 above).
It should be noted that in only six of the 18 Telemachus-yielding TCs are
the bracts sufficiently preserved to allow the execution of line drawings;
and even in these instances, only one or two specimens show a few bracts
that are reasonably complete.
The six Molteno species are based on the following TCs and reference
palaeodemes. All derive from Heidiphyllum thicket, but they are well scat-
tered through the Molteno stratigraphic sequence.
T. elongatus— Tel 111 Hei elo (Telemachus Spruit). 40 indivs
Heidiphyllum thicket; Cycle 3 (Mayaputi Member)
T. grandis— Kan 112 Hei elo (Kannaskop), 10 indivs
Heidiphyllum thicket; Cycle 3 (Mayaputi Member)
T. serribractus— Kle 111 Hei elo (Kleinhoek), 25 indivs
Heidiphyllum thicket; Cycle 2f (Indwe Member)
T. brachybractus — Pen 411 Hei elo (Peninsula), 17 indivs
Heidiphyllum thicket; Cycle 2f (Indwe Member)
T. dubibractus— Qac 111 Hei/Dic (Qachasnek), 10 indivs
Heidiphyllum thicket; Cycle 2c (Indwe Member)
T. acutisquamus— Aas 611 Hei elo (Aasvoelberg), 6 indivs
Heidiphyllum thicket; Cycle 1 (Bamboesberg Member)
Uncertainty— These six species are recognised by JMA. but with
reservation by HMA who feels that there is no bract preserved in the Qac
111 specimens (T. dubibractus ) and agrees that although the scale lobes
appear distinct in the Aas 611 form (T. acutisquamus). this could be due to
some difference in preservation. Furthermore, HMA does not recognise the
bract (as drawn) in the PRE/F/15316 specimen (tf. 9 above) of this Aas 611
palaeodeme.
VOLTZIALES
Telemachus
86
(§f TREL ITZIA 15 (2003)
Telemachus elongatus H.M.And. 1978
Holotype
Specimen: BP/2/5637a,b (C-T.S.326a,b; H.M.And. 1978); pi. 11(1-3).
Assemblage (TC): Tel 111 Hei elo; Telemachus Spruit.
Preservation: virtually complete cone, part and counterpart, longitudinal
section; compression with ferruginised woody remains, ca 60% flat-
tened; impression in thickly laminated, light olive-grey shale with poor
cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 40 indivs (7 intact, 23 partial, 10 detached scales),
tfs 1-4 adjacent, pi. 11(1-12).
Sister palaeodemes— 4 (best 3 listed)
Gre 121 Hei elo: >100 indivs (90 partial).
Aas 111 Hei elo: 12 indivs (1 intact, 3 partial).
Lut 111 Hei/Dic: 2 indivs (1 intact, 1 detached scale).
Specific diagnosis
A Telemachus cone bearing long (ca 23 mm), narrowly lanceolate,
gradually tapering bracts with finely dentate proximal margins.
Specific characters
Strobilus: of intermediate size (50 mm long).
Megasporophyll: bract 23 mm long, narrowly lanceolate, tapering gradu-
ally to acute tip, proximal margin finely dentate.
Etymology
elongatus (Lat.) — with reference to the elongate bracts.
Comment & comparison
Identified from five Molteno TCs, this appears to be the most fre-
quently occurring of the Telemachus species. It is characterised by the par-
ticularly long, finely toothed bracts.
Telemachus grandis J.M.And. & H.M.And , sp. nov.
Holotype
Specimen: PRE/F/20080a,b; pi. 12(1-3).
Assemblage (TC): Kan 112 Hei elo; Kannaskop.
Preservation: virtually complete cone attached to stout pedicel, part and
counterpart, longitudinal section, with a few in situ seeds; impression
in very thin-bedded, moderately baked, medium grey shale with poor
cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 10 indivs (1 intact, 1 partial, 8 detached scales),
tf. 6 adjacent, pi. 12 (1-5).
Sister palaeodemes— nil.
Specific diagnosis
A Telemachus cone of particularly large size bearing intermediate-
length (ca 13 mm) broadly lanceolate bracts without evident dentition
along proximal margins.
Specific characters
Strobilus: large (ca 70 mm long).
Megasporophylls: bract ca 13 mm long, broadly lanceolate, with no appar-
ent marginal dentition.
Etymology
grandis (Lat.)— with reference to the relatively large size of the cones.
Comment & comparison
This species is unique to Kan 112, interpreted as a crevasse-splay
deposit. The cone is distinctly larger than in the other Molteno species.
Telemachus serribractus J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: BP/2/589 la,b; pi. 13(1-3,6).
Assemblage (TC): Kle 1 1 1 Hei elo; Kleinhoek.
Preservation: complete cross-section of cone, part and counterpart, with a
few in situ seeds, impression in very thin-bedded, medium light grey
shale with moderate cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 25 indivs (3 intact, 5 partial, 17 detached scales),
tf. 7 above, pis 12(1-7), 13(1M).
Sister palaeodemes— 1 only.
Mat 111 Die dub: 12 indivs (3 intact, 7 partial).
Specific diagnosis
A Telemachus cone bearing intermediate-length (ca 17 mm) lanceolate
bracts with pronounced proximal shoulders and marked dentition.
Specific characters
Strobilus: of intermediate size (ca 50 mm long).
Megasporophyll: bract ca 17 mm long, lanceolate, tapering relatively
abruptly, proximal shoulders pronounced and strongly dentate.
Etymology
serribractus (Lat.)— with reference to the serrate bracts.
Comment & comparison
This species is known from two TCs, Kle 1 1 1 and Mat 111. Its recog-
nition, as distinct from T. elongatus , seems justified on the basis of the
more abruptly tapering, strongly dentate bracts.
Telemachus
VOLTZIALES
TRELITZIA 15 (2003)
87
Telemachus brachybractus J.M.And. & H.M.And., sp nov.
Holotype
Specimen-. PRE/F/18070; pi. 15(1).
Assemblage (TC): Pen 411 Hei elo; Peninsula.
Preservation : half-complete cone, without counterpart; longitudinal sec-
tion; seeds dehisced; impression in thickly laminated, greenish grey
shale with moderate cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 50 indivs ( 10 intact, 30 partial, 10 detached scales),
tfs 1-3 adjacent, pi. 15(1-9).
Sister palaeodemes— 1 only.
Pen 311 Hei elo; 17 indivs (10 partial).
Specific diagnosis
A Telemachus cone of particularly small size bearing short ( ca 7 mm)
lanceolate bracts with strongly pronounced proximal shoulders and marked
dentition.
Specific characters
Strobilus: small (25-35 mm long).
Megasporophyll : bract ca 7 mm long, lanceolate, tapering fairly abruptly,
proximal shoulders pronounced and strongly dentate.
Etymology
brachybractus (Lat.)— with reference to the short bracts.
Comment & comparison
This species is represented by only two palaeodemes from TCs
(Heidiphyllum thicket of the floodplain) occurring within the same inter-
mittently outcropping mudstone horizon (50-250 mm thick) some 2 km
apart along an unpaved road. The cones, of which there are a good number
of intact specimens, are consistently smaller than in the other five Molteno
species. The small, stout scales are quite distinctive.
Telemachus dubibractus J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/1703a,b; pi. 16(7).
Assemblage (TC): Qac 111 Hei/Dic; Quachasnek.
Preservation: intact cone (proximal third missing off edge of slab), with
part and counterpart; longitudinal section; seeds apparently shed;
impression in thinly laminated, medium light grey shale with moder-
ate cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 10 indivs (3 intact, 2 partial, 5 detached scales), tfs 6-9 adja-
cent, pi. 16(7-10).
Sister palaeodemes — nil.
Specific diagnosis
A Telemachus cone (apparently) bearing very short (ca 4 mm) conical
bracts without proximal shoulders or serrations.
Specific characters
Strobilus: of intermediate size (? ca 50 mm long).
Megasporophyll: bract ca 4 mm long, conical, apparently without proxi-
mal shoulders or serrations.
Etymology
dubibractus (Lat.)— with reference to the dubious nature of the bracts.
Comment & comparison
T. dubibractus is unique to Qac 111. This species is provisionally dif-
ferentiated on the basis of the short bract which is not readily distinguished
from the scale lobes. It is assumed (uncertainly) that the bracts are pre-
served intact (have not been dehisced) in the type specimen— as is the con-
dition of preservation in the many more or less complete cones represent-
ing the other Molteno species.
Telemachus acutisquamus j.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/15318a,b, pi. 16(11).
Assemblage (TC): Aas 611 Hei elo; Aasvoelberg
Preservation: partial cone bearing 6 scales, with part and counterpart;
seeds shed; impression in medium-bedded, strongly baked, dusky yel-
low mudstone with poor cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 6 indivs (2 partial, 4 detached scales),
tfs 10, 11 above, pi. 16(11, 12).
Sister palaeodemes— nil.
Specific diagnosis
A Telemachus cone bearing particularly distinctive scales with gracile,
acutely linear lobes, and with intermediate-length lanceolate bracts of
uncertain proximal character.
Specific characters
Strobilus: unknown.
Megasporophyll: scale lobes gracile, acutely linear; bract ca 9 mm long,
shape and margin uncertain.
Etymology
acutisquamus (Lat.)— with reference to the needle-shaped scales.
Comment & comparison
T. acutisquamus is unique to Aas 611. The remarkably needle-like
scale lobes are very unlike those in the other five Molteno species.
VOLTZIALES
Telemachus
88
d ~f TRELITZIA 15 (2003)
Odyssianthus J.M.And. & H.M.And., gen. nov.
Type species
Odyssianthus crenulatus J.M.And. & H.M.And., sp. nov.
Generic diagnosis
A voltzialean male cone of medium size ( ca 70 mm long), with strong-
ly upcurving microsporophylls bearing 2 rows of ca 4 latero-abaxial clus-
ters (2-3 sacs each) of microsporangia.
Generic characters
Strobilus : a compact cone, lanceolate, relatively large (70 mm long); axis
stout (ca 6 mm in diam.), strongly flexed towards base; microsporo-
phylls helically attached, in ca 23 gyres of ca 1 6-20 units.
Microsporophyll: a simple scale, proportionately intermediate, strongly
upward-curving; distal lamina broadly ovate, margin finely crenulate.
with moderate heel; stalk broad, tapering proximally, winged and
keeled; microsporangia relatively numerous (ca 22-24), latero-abaxial
in two rows of ca 4 pendent clusters, each with 2 or 3 sessile pollen
sacs.
Microsporangium: irregularly rhomboidal (ca 1 mm long), with clear lon-
gitudinal line (dehiscence slit?) and apical ‘micropyle’.
Pollen: unknown.
Eponymy
Odyssianthus— Greek mythology; for Odysseus, the Greek hero at the
siege of Troy and father of Telemachus (the name given to the affiliat-
ed female cone).
Global range: 1 sp., Gondwana, Tr. (CRN).
First & last: the single Molteno species described here.
Gondwana Triassic occurrence
SAf— Karoo Basin, 1 TC (2 indivs).
Molteno occurrence
Frequency (F): 1 TC.
Diversity (D); 1 species.
Abundance (A): 2 individuals total.
Tel 111: 2 indivs in 90 man-hrs cleaving (1 per 5 man-days) extremely rare
The extreme rarity of the male cone is remarkable considering the fre-
quency and abundance of the affiliated foliage Heidiphyllum and the wide-
spread occurrence of the female cone Telemachus (see pp. 82, 83).
Affiliated organs
Female strobilus: Telemachus— Grade 4 (Kin. reinf., Mut. occ.).
Foliage: Heidiphyllum— Grade 4 (Kin. reinf.. Mut. occ.).
The affiliation of Odyssianthus with the ovulate cone Telemachus and
the foliage genus Heidiphyllum is considered sure (Grade 4 reliability) for
reasons of co-occurrence and morphological similarity.
Classification & comparison
Suprageneric classification (Voltziaceae/Voltziales)
Odyssianthus, in view of its Grade 4 affiliation with Telemachus
(female cone) and Heidiphyllum (foliage), rather than its morphology, is
included in the Voltziaceae. Telemachus (And. & And. 1989, pp. 420-423;
this vol., p. 82), in particular, is a typical member of the family.
Morphologically, with its winged scale and double row of latero-abax-
ial microsporangia, Odyssianthus is very distinct from the Laurasian gen-
era— Sertostrobus, Darneya and Willsiostrobus, best known from the lower
Middle Triassic Gvcs-n-Voltzia Sandstone of France (Grauvogel-Stamm
1978) — that are generally included in the family (this vol., pp. 57, 59).
Considering a possible phylogeny based exclusively on male cones (Tab.
31, p. 57), Odyssianthus (plus Lutanthus) could quite readily be derived
from the Permian glossopterid genus Lidgettonia. This option would place
Odyssianthus well distant from the Voltziales, in a new gymnospermous
order and would seemingly negate affiliation with Telemachus.
Intergeneric comparison (Molteno genera)
While Lutanthus and Fredianthus are included, possibly conservative-
ly, along with Odyssianthus in the Voltziales (an order transitional between
the Cordaitanthales and Pinales), they might well be shown, once their ovu-
late and foliage affiliates are known, to represent different lineages at order
level.
Reconstruction
The two available specimens of Odyssianthus are excellently pre-
served, clearly showing both the outer aspect of the cone and longitudinal
sections revealing the characteristics of the scales and microsporangial
clusters. Even so, the precise number of microsporangial groups and
microsporangia per group is not as certain. The base, also, is not clear, but
it appears that the cone, as in Lutanthus, is sessile (without a free axis).
Adaptive radiation (Molteno diversity)
With Odyssianthus sampled from only the single site, it is unknown
how morphologically varied or conservative the male cone of this plant-
genus might have been with respect to the female cone (Telemachus) or
foliage ( Heidiphyllum ). Would the pollen cones, as in the case of
Rissikianthus (pp. 108-1 1 1 ), reflect the same level of species diversity for
the genus as the ovulate cones do? The possibility does exist, though we
find this unlikely, that either or both of the male cone genera, Fredianthus
and Lutanthus (pp. 70, 71, 74-77), might be congeneric with Odyssianthus.
If so, the diversity shown by the male cones of the Telemachus! Heidi-
phyllum plant-genus would be particularly unusual.
Affiliated organs (elaborated)
Mutual occurrence: As with the other voltzialean male cones from the
Molteno, Odyssianthus remains extremely rare. It is known only from the
two intact specimens from Telemachus Spruit (Tel 111), where
Heidiphyllum occurs as a monodominant (89% of the foliage in the assem-
blage) and Telemachus is relatively numerous (40 specimens). No other
coniferous remains, either foliage or cone, are known from this Tel 1 1 1 TC.
Morphological correspondence: In both size and texture Odyssianthus and
Telemachus are very alike, but the one is clearly microsporangiate and the
other ovulate.
Odyssianthus
VOLTZIALES
d^TRELITZIA 15 (2003)
89
Odyssianthus crenulatus j.M.And. & H.M.And., sp. nov.
Holotype
Specimen. PRE/F/17368a,b; pi. 17(1-7).
Assemblage (TC): Tel 111 Hei elo; Telemachus Spruit.
Preservation: almost complete intact cone (free axis missing), part and
counterpart, longitudinal section and outer view, showing numerous
scales and in situ microsporangia; impression in thickly laminated,
light olive-grey shale with poor cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 2 individuals, including the holotype and a second fairly com-
plete cone (PRE/F/18286a,b; pis 17(8, 9), 18(1-7).
Sister palaeodemes— nil.
Specific diagnosis— as for genus.
Specific characters — as for genus.
Etymology
crenulatus (Lat.)— with reference to the crenulate nature of the scale margin.
Comment & comparison— see notes for genus.
(). crenulatus
Tel 1 1 1
GONDWANA TRIASSIC, GEOSTRAT’ DISTRIBUTION
VOLTZIALES
Odyssianthus
90
d/TRELITZIA 15 (2003)
Heidiphyllum Retallack 1981b
Type species
Heidiphyllum elongatum (Morris 1845) Retallack 1981b.
‘Jerusalem Basin’. Tasmania; U. Triassic.
Generic diagnosis
A voltzialean leaf of relatively large size, with ca 10 parallel veins,
attached to short shoots in tight pseudowhorls with short triangular bracts.
Generic characters
Attachment: leaves in tight pseudowhorls on short shoots, in axils of per-
sistent triangular bracts.
Leaf, individually dehisced, relatively large (ca 120 X 10 mm), blade
broad, flattened, simple, linear to narrowly elliptic or oblanceolate,
apex rounded, base narrowly to broadly sessile; veins parallel, moder-
ately spaced (ca 10 per 10 mm), forking nearer base, coalescing to-
wards apex.
Cuticle (adapted from And. & And. 1989, p. 429); this vol., tfs 4-6 below.
Yield: Lit 1 1 1 : 13 specimens sampled; grade index 0/1 1/2/0/0.
Umk 111: 19 specimens sampled; grade index 1/7/1/0/0.
Diagnostic characters: based on H. elongatum (Umk 111).
Adaxial/ abaxial: dorsiventral, U & L cuticle of equal thickness.
Cell characteristics: walls straight to gently curved, profile triangular, plan
normal; upper cuticle with isodiametric to oblong cells, end walls
square to oblique, 350 cells per 1 mm2, nonpapillate; lower cuticle with
pentagonal to hexagonal cells, 600 cells per 1 mm2, papillae large,
bold, usually 2 per cell.
Stomatal apparatus: haplocheilic, ?amphistomatic; subsidiary cells ano-
mocytic, 5 or 6 cells, noncutinised, radial walls normal; Florin ring
lappetate, interfingering baculae; (frequency, arrangement, guard cells
and stomatal pit unknown).
Other features: none preserved.
Eponymy
Heidiphyllum —named by Retallack (1981b) after Heidi M. Anderson who
first described the affiliated cones (Anderson 1978).
Global range: 3 spp., Gondwana, Tr. (ANI-NOR).
First: H. elongatum (Desmiophyllum sp.) (Lele, 1962b); Beli, S. Rewa, India.
Last: H. elongatum (Retallack 1985); Wairoa Gorge, Nelson Synch, N.Z.
Gondwana Triassic occurrence
Frequency (F): 26 degree squares (of 84 across Gondwana).
Ubiquity (U): 5 continents (of 5 comprising Gondwana).
Diversity (D): 3 foliage species.
Abundance (A): 95% (a monodominant in Molteno TCs).
Longevity (L): 18 myrs (late Anisian to late Norian).
Colonisation success: FUDAL rating 26/5/3/95/18 = 147.
High success (Grade 2): Heidiphyllum is the second most prominent
genus in the Gondwana Triassic; it was frequent, ubiquitous, abundant
and long-lived, but markedly lacking in diversity.
Endemism: of the three Gondwana Triassic species, one is a basin endem-
ic, one a continent endemic, while the third (H. elongatum ) occurs very
widespread throughout the realm.
Molteno occurrence
Frequency (F): 62 TCs (of 100 sampled in the Molteno).
Diversity (D): 1 species.
Abundance (A): monodominant (>70%) in its preferred habitat at 19 TCs;
co-dominant (20-69%) at 14 TCs; occasional to abundant (1-19%) at
24 TCs; <1% at only 5 TCs.
Habit: probably a woody, reed-like plant.
Preferred habitat: Heidiphyllum is monodominant in areas of high water
table in the floodplain or on channel sandbars.
Affiliated organs
Female strobilus: Telemachus— Grade 4 (Kin. reinf., Mut. occ.).
Male strobilus: Odyssianthus— Grade 4 (Kin. Reinf., Mut. occ.).
Classification & comparison
Gondwana Triassic: Axsmith et al. (1998b) describe Notophyton
krauselii , a permineralised leaf from the Triassic of Antarctica, and consid-
er it as possibly conspecific with H. elongatum. They record the leaves as
being attached either helically or bijugately (opposite), which would make
them more similar to Clariphyllum. They find their material to have a pos-
sible link to the Podocarpaceae.
Other attached leaves are recorded by Zhou & Zhang (1998) for
Phoenicopsis euthyphylla from the Middle Jurassic Yima Fm., China.
These are attached to short shoots which are in turn attached to a long
shoot. Associated with these leaves is Tianshia patens (similar to
Clariphyllum), a leafy shoot with helically arranged leaves, which the
authors suggest belongs to the same plant as P. euthyphylla.
Heidiphyllum
VOLTZIALES
G^TRELITZIA 15 (2003)
91
3
Reconstructions
The reconstruction of the leaves (tfs 1,2 adjacent) in pseudowhorls on
short shoots attached to an axis is based on a series of specimens from Aas
311: PRE/F/22063 (tf. 2 opposite) shows an axis bearing a small short
shoot with leaves attached, probably in a whorl (on this specimen more
short shoots and leaves occur but not so clearly); PRE/F/22061 (tf. 3 oppo-
site) and PRE/F/22066 show leaves attached to axes with remains of leaf
bases and triangular bracts; PRE/F/1 1 894 shows attached triangular scales.
We have used the extant conifer Pseudolarix amabilis from China as an
anologue. In this taxon the leaves are deciduous and borne on short shoots
in a tight spiral. Another extant tree with leaves (in this case evergreen) on
short shoots in such a tight spiral that they look whorled is Sciadopitys ver-
ticil lata from Japan.
The occurrence of short shoots and fairly wide stems points to
Heidiphyllum being woody and not herbaceous like Aetophyllum discussed
below. Whether Heidiphyllum leaves were deciduous or evergreen is
unknown, but we regard them as having probably been evergreen.
Literature (supporting evidence & viewpoints)
Retallack (1977)— Australia, Clarence-Moreton Basin, Nymboida Colliery,
Middle Triassic (Ladinian); Heidiphyllum found as a very dominant leaf in
the lower part of crevasse splay sandstones, suggesting that it formed a
‘levee and point bar scrub or woodland whose leaf litter was scoured out
during floods’.
Grauvogel-Stamm (1978)— France, Vosges, Gres a Voltzia , Buntsandstein,
late Lower Triassic (Spathian); Aetophyllum, from the Vosges, must be the
most completely known genus within the Voltziales. The material includes:
10 seedlings (five nearly complete) with roots, young stems and leaves;
mature plants (one complete male, six nearly complete) with either female
or male cones found attached, suggesting plants of 2 m height. All speci-
mens are found near the base of fossiliferous lenses, and judged to be pre-
served close to their position of growth. Since only a small amount of
wood— i.e. secondary xylem— is found in the stems studied, Aetophyllum
is considered by Grauvogel-Stamm to be herbaceous, a condition previ-
ously unknown amongst the Pinopsida. The interpretation is supported by
Roth well et al. (2000).
Meyer- Berthaud & Taylor (1991)— Antarctica, Fremouw Fm., Silicified
peat locality, Fremouw Peak.
Adaptive radiation (Molteno diversity)
Though Heidiphyllum is frequent, occurring in 62 TCs (Tab. 44) and
abundant (often monodominant), it is morphologically particularly conser-
vative. Only one apparent species of foliage can be recognised in this great
mass of material from the Molteno. The distinct species from Umk 111.
previously recognised by us as a species of Heidiphyllum, is here placed in
the new genus Clariphyllum (pp. 100, 101). Two additional species are
recognised (And. & And. 1989) in collections from Gondwana deposits
further afield.
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
1-10: degr!
O superb
e squares j
^alities j
2
ir‘"
envisage
erect snr
tree with
branche
d as a
ub to s
simplt
arge
nail
side
&
4 '-Qy
5
/ ■ • 6
7
c
8 W f
i
9
r oi
10
Molt
E
LL
O
C
VOLTZIALES
Heidiphyllum
92
^TRELITZIA 15 (2003)
BP/2/5637b
BP/2/5637b
Hoiotype Telemachus Spruit
(Tel 111 Hei elo)
PRE/F/7693b
v e
. . . ■- 5SK
, * mel i u 1
3 PRE/F/7693b
' BP/2/5645
PRE/F/17371
BP/2/5642
BP/2/5642
BP/2/5645 ’•
PRE/F/17370
PRE/F/7692
pi. 11
Telemachus elongatus
VOLTZIALES
93
c/tRELITZIA 15 (2003)
PRE/F/20080b
? BP/2/3415
T. elongatus
PRE/F/20080a
Aasvoelberg
(Aas 1 1 1 Hei elo)
T. elongatus
X4 > ' BP/2/4326
BP/2/4328
PRE/F/19440
VOLTZIALES
pi. 12
Telemachus spp.
df TRELITZIA 15 (2003)
BP/2/5891a
Holotype
Kleinhoek
(Kle 1 1 1 Hei elo)
Telemachus serribractus
pi. 13
VOLTZIALES
Ci/TRELITZIA 15 (2003)
95
Matatiele
(Mat 111 Die dub)
v Kleinhoek
' (Kle 1 1 1 Hei elo)
PRE/F/9264a
PRE/F/9264b
T. serribractus
pi. 14
VOLTZIALES
Telemachus serribractus
96
df' TRELITZIA 15 (2003)
PRE/F/18070
Holotype
Peninsula
(Pen 411 Hei elo)
PRE/F/17177a
SHE
PRE/F/17180
Si PRE/F/18068
PRE/F/17180
PRE/F/17211
PRE/F/18055b
PRE/F/18052
PRE/F/ 19632
Telemachus brachybractus
pi. 15
VOLTZIALES
97
c/tRELITZIA 15 (2003)
Peninsula
(Pen 3 1 1 Hei elo)
T. brachybractus
Qachasnek
(Qac 1 1 1 Hei/Dic)
T. dubibractus
Aasvoelberg
(Aas 61 1 Hei elo)
T. acutisquamus
PRE/F/16949a
PRE/F/16941a
pi. 16
Telemachus spp.
VOLTZIALES
98
^VtRELITZIA 15 (2003)
WSMtr
PRE/F/ 17368c
PRE/F/17368C
\%V.
PRE/F/17368b
PRE/F/17368a Holotype
PRE/F/18286b ... <B
PRE/F/18286a
Telemachus Spruit
(Tel 1 1 1 Hei elo)
PRE/F/17368a
; X5 ' PRE/F/17368a
PRE/F/17368a
Odyssianthus crenulatus
pi. 17
VOLTZIALES
99
^'/trELITZIA 15 (2003)
_/ -jt
all PRE/F/18286a
Telemachus Spruit
(Tel 111 Hei elo)
X40
VOLTZIALES
pi. 18
Odyssianthus crenulatus
100
<§?'. TRELITZIA 15 (2003)
Clariphyllum J.M.And. & H.M.And., gen. nov.
Type species
Clariphyllum clarifolium J.M.And. & H.M.And. 1989, comb. nov.
Generic diagnosis
A voltzialean leaf of relatively small size, with ca 4 parallel veins,
attached to shoots in a lax spiral arrangement without bracts.
Generic characters
Attachment-, leaves in lax spiral, on long shoot, without bracts.
Leaf, individually dehisced, leaf small (75 X 3.5 mm), linear elliptic; tip
rounded obtuse, base narrowly sessile; veins parallel, moderately
spaced (10 per 10 mm), consistently at midlength, once forked near
base, coalescing towards apex.
Cuticle : this vol., p. 101.
Eponymy
Clariphyllum'. Clara’s leaf, in honour of our eldest daughter who accom-
panied us on several collecting trips.
Global range: 1 sp.. Gondwana, Tr. (CRN).
First & last: the single Molteno species described here.
Gondwana Triassic occurrence
Frequency (F): 3 degree squares (of the 84 across Gondwana).
Ubiquity (U): 1 continent (of 5 comprising Gondwana).
Diversity (D): 1 foliage species.
Abundance(A): <1% (the norm in Molteno TCs).
Longevity (L): 1 myrs (Camian).
Colonisation success: FUDAL rating 3/1/1/- / 1 =6.
Minimum success (Grade 1): Clariphyllum was the 24th most promi-
nent genus in the Gondwana Triassic; it was very low in frequency,
ubiquity, diversity, abundance and longevity.
Endemism: the single species is endemic to the Molteno.
Molteno occurrence
Frequency (F): 3 TCs (of 100 sampled in Molteno).
Diversity (D): 1 species.
Abundance (A): 51 individuals total, rare to extremely rare.
Habit: probably a shrub.
Preferred habitat: Dicroidium riparian forest.
Affiliated organs
Female strobUus: unknown.
Male strobilus: unknown.
Classification & comparison
Intergeneric comparisons
Gondwana Triassic pinopsid genera— Clariphyllum was originally
included by us in Heidiphyllum (And. & And. 1989), but considering the
very different mode of attachment, the narrow leaflets and reduced number
of veins, and the seemingly different cuticular preservation, this taxon is
described here as a distinct genus. It is placed in the Voltziales on the gen-
eral appearance of the individual leaves, but without any clue as to its affil-
iated female or male cones, or evidence of cuticular morphology, this can-
not be verified.
Non-Gondwana genera— For discussion and comparison with
Tianshia patens from the Middle Jurassic of China, see text for Heidi-
phyllum (p. 90).
Clariphyllum
VOLTZIALES
C?/tRELITZIA 15 (2003)
101
Clariphyllum clarifolium j.M.And. & H.M.And. 1989,
comb. nov.
Holotype
Specimen: PRE/F/6582 a,b; And. & And. [1989, pi. 264(10-12)].
Assemblage (TC): Umk 111 Die 2spp; Umkomaas Valley.
Preservation: compression, a foliage shoot with ca 10 leaves, part and counter-
part.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 20 indivs. And. & And. [1989, pi. 264(1-17)].
Sister palaeodemes— 2 only.
Maz 211: 1 indiv.
Kap 111: 30 indivs.
Specific diagnosis— as for genus.
Specific characters— as for genus.
Eponymy
clariphyllum— for our daughter Clara who has accompanied us on several
collecting trips.
Comment & comparison
At Kap 111 and Maz 211 there have been found no attached leaves of
this form and thus their generic identity needs future confirmation.
Cuticles
Potential sample: Lit 1 1 1 , 20 indivs.
Macerated (this work): 9 indivs.
Presen’ation grade: Grade 1 .
Diagnostic characters: none obtained.
Comment: —
Significance: of no value in classification and affiliation.
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
Ileidiphyllum
II. minutifolium
II. cacheutense
redrawn after Walkom, 1924, pi. 21 (3A),
Bellevue Station, Esk Fm.(Laainian)
Clarence/Moreton Basin,
Queensland
These two narrow-leaved species of
Heidiphyllum — as recognized in
And, & And,, 1989 — are included
here for direct comparison with C.
clarifolium. Though both the South
American and Australian species
have leaves about as narrow as C.
clarifolium, they show a quite differ-
entpattem ofvenation.
redrawn after Kurtz, 1921 , f 22(330,323),
Estratos con Estheria, Cacheuta Fm. (Camian),
Cacheuta Subregion,
Argentina
tfs 1-3
from And. &And.,
VOLTZIALES
Clariphyllum
102
d/TRELITZIA 15 (2003)
PINOPSIDA S.V.Meyen 1984
PINALES (Coniferales)
PODOCARPACEAE Endl 1847
Rissikistrobus J.M.And. & H.M.And., gen. nov.
Type species
Rissikistrobus plenus J.M.And. & H.M.And., sp. nov.
Generic diagnosis
A pinalean female cone of linear shape with bract/scale complexes of
1-3 lobes bearing a pair of adaxial ovules on each lobe.
Generic characters
Attachment : terminal on leafy axis.
Strobilus: cone compact, linear (to ca 100 X 5 mm), megasporophylls spi-
rally arranged, ca 6 units per gyre.
Megasporophyll: cone units consisting of ovuliferous bract/scale com-
plexes; bracts leafy (to 5 mm), lanceolate, cuspidate, erect; scales, 1-3
lobed; ovules adaxial, paired, in concave surface of lobe.
Ovule : naked, (?)spathulate, 1.5 mm long.
Etymology
Rissikistrobus— after the affiliated foliage Rissikia.
Global range: 3 spp., Gondwana, Tr. (? - CRN).
First: Rissikistrobus semireductus (Retallack 1977); Cloughers Creek Fm.
(UNEL 1564), Nymboida, Australia.
Last: the 3 Molteno species described here.
Gondwana Triassic occurrence
SAf— Karoo Basin, 8 TCs ( ca 80 indivs).
Aus— Clarence Moreton Basin, 2 TCs (?indivs).
Although the affiliated foliage Rissikia occurs widespread in the
Gondwana Triassic, Rissikistrobus (a single small fragment) has only been
recorded from the Cloughers Creek Fm., Nymboida CM (Retallack et al.
1977) and according to K. Holmes (pers. comm.) also occurs in his collec-
tions from Basin Creek Fm., Nymboida CM Australia.
Molteno occurrence
Frequency (F): 7 TCs (of 100 sampled in the Molteno).
Diversity (D): 3 species.
Abundance (A): 85 individuals total, very rare to extremely rare.
Hla 212 Die 3spp:
2 indivs in
11
Umk 1 1 1 Die 2 spp:
45 ” ”
400
Hla 213 Die elo:
5 ” ”
60
Kon 222 Die edo:
2 ” ”
40
Aas 411 Dic/Sph:
20 ” ”
512
Kap 111 Dic/Ris:
3 ” ”
65
Lit 111 Dic/Hei:
8 ” ”
550
in 1 1 man-hrs cleaving (2 per 1 man-days) very rare
) extr. rare
For details of the frequency and abundance of the three affiliated
organs, Rissikistrobus (female), Rissikianthus (male) and Rissikia (female),
through the Molteno, see Tab. 38 (p. 104); and for analysis of the mother
plant’s preferred habitat and of a typical TC (Pen 321 Dic/Ris) in which it
occurs as a co-dominant, see And. & And. (in prep.).
Affiliated organs
Female: Rissikistrobus— Grade 4 (Cut. cor., Mut. occ.).
Foliage: Rissikia— Grade 4 (Cut. cor., Mut. occ.).
The affiliation of fruit (male and female) and foliage, based on the dis-
tribution data tabulated for the Molteno (Tab. 38, p. 104), is virtually cer-
tain. Townrow (1967) established to his satisfaction that the cuticle of the
three organs corresponded. We have not attempted to verify his results
based on specimens from our collection (see further on pp. 104, 105).
Classification & comparison
Suprageneric classification (Podocarpaceae/Pinales)
It might be expected that within the initial Triassic radiation of the
Pinales (pp. 56, 57), during which six of the eight extant families are gen-
erally accepted to have emerged, there would be encountered many archa-
ic and transitional features within the evolving lineages. This seems to be
well exemplified, for instance, in the Rissikistrobus/ Rissikia plant, where
characters of the sister families Pinaceae and Podocarpaceae are both in
evidence. Considerable published debate (p. 105) has been devoted to
which of the two families is represented by this widespread Gondwana
Triassic genus (see map for Rissikia, p. 1 13). The debate is not settled and
we have wavered this way and that, settling, for now, with no great con-
viction on the Podocarpaceae.
Reconstructions
For each of the three species, R. plenus, R. semireductus and R. reduc-
tus, an R4 grade reconstruction has been made and each is based on one or
two particular specimens along with the rest of the reference palaeodeme
for the taxon.
The precise nature of the scale/bract complexes in the three recognised
species of Rissikistrobus has not been unambiguously established— as is
demonstrated in the degree to which we differ from the morphological
interpretations of Townrow (1967) outlined on p. 105. Our various recon-
structions of both the megasporophylls and the full cones should be accept-
ed in this light. No doubt more cuticular studies of the reasonably numer-
ous Umkomaas (Umk 111) and Little Switzerland (Lit 111) specimens
together with sectioning of the casts/moulds of the Aasvoelberg (Aas 411)
specimens, should go a long way towards clarifying the structure.
Rissikistrobus
PINALES
c/trelitzia 15 (2003)
103
Molteno occurrence (elaborated)
Mother plant ( RissikistrobusIRissikialRissikianthus )
The habit and preferred habitat of the mother plant of Rissikia media,
by far the more frequent and abundant of the two Molteno foliage
species, is analysed in some detail in a sequel to this work (And. & And.,
in prep.). It is visualised as being a substantial, scattered tree in Dicroi-
diutn riparian forest or Dicroidium open woodland, occasionally forming
monodominant wetland stands.
Foliage ( Rissikia )
Rissikia occurs in 21 of the 100 Molteno TCs and in all but one
instance in assemblages dominated by Dicroidium— in riparian forest and
open woodland. Generally the genus is rare (<1%), but in five TCs it does
become fairly common (1-5%) and in two TCs it is a co-dominant
(3CM-0%). In the latter two instances, Rissikia is seen as forming a dis-
tinct vegetation type — akin to the swamp Cyprus — within the open wood-
land.
Female cone ( Rissikistrobus )
The female cone ( Rissikistrobus ) occurs in eight TCs, twice as often
as the male. It clearly appears more often in the riparian forest TCs than
elsewhere. Even where the male and female occur in the same assem-
blage, their abundance tends to be very unbalanced, either the one way or
the other.
Male cone ( Rissikianthus )
The male cone ( Rissikianthus ) occurs in only five TCs: essentially
once in each vegetation type— and in each case where the foliage is most
common for that habitat. The conclusion drawn from this is that the male
occurs only autochthonously or very nearly so. Interestingly, in the two
instances (Kap 111 & Pen 321) where the foliage is particularly abundant,
the male is clearly more common than the female.
Gondwana Triassic occurrence (elaborated)
Whole-plant genus {Rissikistrobus! Rissikia! Rissikianthus)
The whole-plant genus is well established on the basis of the secure
affiliation (Grade 4) of all three organs — foliage, female cone and male
cone. While the ‘genus’, through its foliage, is known to have occurred
widespread across Gondwana, the reproductive organs remain virtually
unknown beyond the Molteno.
Foliage genus (Rissikia, pp. 112, 113)
Rissikia has been established in this volume (Tab. 20, p. 27) as the
11th most prominent of the 27 Molteno gymnospermous foliage genera
in the Gondwana Triassic. With a FUDAL rating (colonisation success)
of 39, it occurred ubiquitously across Gondwana during the upper half of
the Triassic, being recorded from no less than 1 1 degree squares outside
‘Africa’. Its abundance is not recorded in the literature, but it probably
occurred only as an occasional element in most TCs (1-2%) as in the
Molteno; and it reflected little diversity (two species).
Ovulate-cone genus ( Rissikistrobus )
In marked contrast to the foliage, Rissikistrobus has been recorded,
aside from Africa, only in eastern Australia: one published and one
unpublished record from the Nymboida Coal Measures (Retallack et al.
1977; Holmes, pers. comm.).
Pollen-cone genus (Rissikianthus, pp. 108, 109)
As for the other four pinopsid male-cone genera recorded in the
Molteno, Rissikianthus remains unknown elsewhere in Gondwana.
Tab. 37.
RISSIKISTROBUS HYPODIGM, Gondwana Triassic occurrence
AUTHOR
SUBREGION
FORMATION
LOCALITY
NAME
Indivs ILLUSTRATION
SOUTH AFRICA
1967jtownrow
Underberg
KA9
24
Molteno
Umkomaas
Rissikia media
2
pi 1 (A.B), tf 8(C-J)
” |
”
” apiculati
1
pi 1(C,D), tf 8(D)
1978-1999 Anderson & Anderson Molteno literature not included in this table
AUSTRALIA
1977 Retal. et al.
Nymboida
CM7
19
Cloughers Cr.
UNEL 1564 (Kang.Cr.)
Rissikia media
1
f 11(A)
Molteno
Species
I
Intact-
ness
s E
2. E “
u o> ™
IS m o
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
PINALES
Rissikistrobus
104
c/tRELITZIA 15 (2003)
assemblages
(taphocoenosis)
Rissikia
O Rissikistrobus
R. pienus
R. semireductus
R. reductus
Incertae
O seeds (in-situ)
Q. Rissikianthus
R. concavus
R. linearis
R. townrowii
R. convectus
Incertae
microsporangia
(in-situ) *
Boe 112 Die cor
12
1
l! - - - - 1 1
Tin 121 Sph 2spp
1
Kon 223 Die odo
4
” 222 " "
9
2] - - - 2 -
” 111 " "
2
-1 -1 -i -( -1 -)
Pen 321 Dic/Ris
35
30 1 30 - - - - 7
" 211 Dic/Equ
1
!
" 221 Dic/Equ
2
- j ; - ; -
" 421 Die odo
5
Kle 111 Hei/Dic
3
- 1 - - - - 1 -
-1
Kap 111 Dic/Ris
38
3! - - - 3 -
25! - - - 25 - 5
Nuw 111 Die zub
2
. . .
Hla 211 Die 3spp
1
1
" 212 Die 3spp
1
2; - 2 - - -
. .1 .1 .1 -1 .
" 213 Die elo
2
5' 3 2 - - 2
-1 ----- -
Umk 111 Die 2spp
5
45! 4 41 - - 4
8l - - 8 - - 2
Mng 111 Die 2spp
2
-! -1 -I -! -
-| -! -; -| -: -
Lit 111 Dic/Hei
40
8| - - - 8 -
Aas 311 Hei elo
1
_ 1 _ ! _> _ i _ ( .
" 411 Dic/Sph
25
20' - - 20 - -
15' - 15 - - - 5
Bam 111 Die dub
1
-! -i -; -1 -| -
Total TCs
Total indivs
21
%
7! 2 3! 1 ! 3 > 2
85! 7 45 20 13 6
1
5! 1 1 1 1 1 5
79! 30 15 8 25 1 20
1 i
* microsporangia clearly evident
Tab. 38. RissikistrobusIRissikia, Molteno occurrence
Evidence for affiliation of organs
Short of organic attachment, the affiliation of foliage ( Rissikia ), seed
cone ( Rissikistrobus ) and pollen cone ( Rissikianthus ) is as securely estab-
lished as can be expected.
Mutual occurrence
Rissikia, a relatively frequent component of the Molteno flora, occurs
in 21 TCs (Tab. 38). Rissikistrobus occurs in seven TCs, invariably in co-
occurrence with Rissikia, while Rissikianthus occurs in five TCs, also
invariably together with the foliage, though in only three of these, along
with the ovulate cone. Overall, in the context of fossil floras, this amounts
to particularly strong evidence for three-way affiliation.
Retallack et al. (1977) records Rissikia leaves from eastern Australia
and a female cone from the same locality (Tab. 37). This is here regarded
as Rissikistrobus sp. indet. and provides further evidence of mutual occur-
rence.
Cuticular correspondence
In support of the evidence based on co-occurrence, the strong similar-
ity between the cuticles of the foliage and the reproductive organs
Rissikistrobus and Rissikianthus (see sketches opposite and on p. 112) pro-
vides convincing proof of affiliation.
Nomenclature (as relates to affiliation)
Holotype: Townrow (1967) originally described the female cone
together with the male counterpart and foliage as Rissikia media. We have
retained his name for the leaves and introduce new names here for the
female and male cones. Townrow never nominated a holotype but did sug-
gest that Bumera Waterfall (i.e. Umk 1 1 1 ) be considered the locus typicus.
Intactness & preservation of cones (Molteno)
In situ seeds
The seeds of Rissikstrobus, unlike those of Telemachus, are nowhere
satisfactorily preserved in the Molteno collection. In only four cones from
Umk 111 and two from Hla 213 are the very small, featureless, spathulate
seeds/ovules seen in situ. In the remainder of the 86 available cones, the
seeds have either not developed to maturity or have been dispersed, or the
material is insufficiently preserved.
Dispersed seeds
The Rissikistrobus seed is very much the same size and shape as the
microsporangium of Stachyopitys sp. A. Only under the microscope, con-
sidering the fingerprint ornamentation of the latter and the featurelessness
of the former, are they distinguishable. We have not scanned slabs from the
Rissikia-yielding TCs microscopically to establish the extent of occurrence
of dispersed Rissikistrobus seeds.
Cuticles
Potential sample: Umk 111, 45 indivs.
Macerated (this work): nil.
Preservation grade: Grade 4—5.
Diagnostic characters: see Townrow (1967); this vol., p. 105.
Comment: as for Rissikianthus (p. 109).
Significance: as for Rissikianthus (p. 109).
Adaptive radiation (Molteno diversity)
Identifying species within the genus is difficult and the conclusions
reached here need to be verified by cuticular studies. Only two reasonably
comprehensive and well-preserved palaeodemes (Umk 111 & Aas 4 1 1 ) are
at hand. Though the affiliated foliage, Rissikia, from the two TCs appears
very alike, suggesting the presence of a single species, the female cones are
significantly different.
As in Telemachus , it is in the scale/bract complexes that the differences
between the species appear. A reduction series seems to be revealed in the
cones from Umk 1 1 1 and Aas 411 —from scales with three deeply divided
equal-sized lobes through forms with strongly reduced lateral lobes, to
those at Aas 4 1 1 (clearly) and at Umk 111 (less clearly) where only the cen-
tral lobe occurs. Three species are provisionally recognised. It is not impos-
sible that the Umkomaas (Umk 111) species are, in fact, members of a mor-
phological or ontogenetic series.
The three Molteno species are based on the following TCs/reference
palaeodemes.
R. p/raus-Umk 1 1 1 Die 2spp (Umkomaas Valley). 4 indivs
Dicroidium riparian forest (mature); Cycle 2b (Indwe Member)
R. semireductus— Umk 111 Die 2spp (Umkomaas Valley), 41 indivs
Dicroidium riparian forest (mature); Cycle 2b (Indwe Member)
R. reductus— Aas 411 Dic/Sph (Aasvoelberg), 20 indivs
Sphenobaiera closed woodland; Cycle 1 (Bamboesberg Member)
Rissikistrobus
PINALES
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105
Townrow (1967) on Rissikistrobus/ Rissikia classification
Affinity with the Podocarpaceae
After an exhaustive comparative study between Rissikia (with its
cone affiliates) and the extant coniferalean families, Townrow (1967)
concluded that this Triassic genus could be referred, at least ‘for the pre-
sent’, to the Podocarpaceae, and that ‘the only family that otherwise
enters the picture is the Pinaceae’. He added that ‘even though this does
involve a slight widening of the limits of the family confusion is
unlikely, for the modification needed is small. The alternative would be
a new family, and there seems no object in this.’ There follows, in
briefest outline, Townrow’s more obvious morphological considerations
supporting his conclusion.
Female cone— a) The spike-like cone of ‘ Rissikia ’ is seen in the
podocarps Dacrydium franklinii, Podocarpus spicatus and P. andinus,
but in no other extant conifer family; b) the 3-partite scale of ‘Rissikia’
is faintly reflected in the reduced, 3-pointed scales of the podocarps D.
franklinii, Michrocachrys and Saxegothea\ c) in Podocarpus spicatus
and P. andinus, a double vascular trace— as ‘possibly’ occur in each
scale lobe of ‘Rissikia’ — proceeds ‘past the chalaza into the further part
of the epimatium’.
Male cone— a) The two pollen sacs (per scale) and disaccate pollen
of ‘ Rissikia ’ are found with regularity in the Podocarpaceae and
Pinaceae only; b) ‘the corpus and saccus shape and ornamentation seem
.... to come close to Dacrydium' . (‘Striae are not known in any living
conifer.’)
Foliage— a) Long and short shoots, as occur in ‘Rissikia’, are found
in only a few living conifers falling in three families —Metasequoia and
Taxodium distinctum (in the Taxodiaceae). Larix and Cetrus (in the
Pinaceae ), and Acmopyle, some Podocarpus species and perhaps
Polypodiopsis (in the Podocarpaceae); b) the bilateral leaves character-
istic of Rissikia ‘are found today only in the Podocarpaceae’; c) ‘the
very thin leaves with four stomatal zones each only one stoma wide can
be seen in young Dacrydium novoguineense’ , while ‘the stomatal details
of Rissikia ’ recall the podocarps Saxegothea and Microstrobos ‘espe-
cially’ (though comparison with Metasequoia in the Taxodiaceae is also
possible).
Townrow (1967) reconstructions of Rissikistrobus
On the basis of his detailed morphological study of six compression
specimens from Umk 111 (collected by himself and now housed in the
Australian Museum, Sydney), Townrow (1967) concluded that the
female cone Rissikistrobus was best included in the extant family
Podocarpaceae. If correct, Rissikistrobus and its affiliates would consti-
tute the first appearance of the family (Cleal 1993; and see pp. 57).
This work of Townrow has been cited frequently since by authors
writing on the fossil conifers (e.g. Miller 1977; Taylor et al. 1987;
Stewart & Rothwell 1993; Taylor & Taylor 1993) with varying degrees
of agreement concerning the podocarpaceous affinities of the material.
Miller (1977), for instance, concluded that Rissikia ‘would probably
have been classified in the Voltziales had it not been for the podocarpa-
ceous foliage and the ease with which we can envision the possible evo-
lution of the known Jurassic forms of the Podocarpaceae from this
genus’.
We follow Townrow’s interpretations with uncertainty. For instance,
Compsostrobus from the Late Triassic Pekin Fm. of North Carolina,
U.S.A., which Taylor et al. (1987) place, with reservations, in the
Pinaceae has features reminiscent of Rissikistrobus.
Rissikia
Female sensu Townrow, 1 967 Male
seed cone, bract
1 Rissikia
sensu Townrow, 1967
tfs 13 p ttj.'/ji ti
redrawn after Townrow, 1967
from And. & And., 1989
R. apiculata
R. media
tfs 4-11
from Townrow, 1967
AM: Australian Museum
PINALES
Rissikistrobus
106
d^TRELITZIA 15 (2003)
Rissikistrobus plenus J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/6774a.b; tf. la,b, adjacent, pi. 19(1-7).
Assemblage (TC): Umk 111 Die 2spp; Umkomaas Valley.
Preservation: incomplete cone (proximal end missing), part and counter-
part, longitudinal outer view showing many scales with in situ ovules;
compression in thinly laminated, carbonaceous (good cuticle) moder-
ately baked, dark grey shale with good cleavage.
Reference palaeodeme
Assemblage: as for holotype.
Specimens: 4 indivs (4 intact cones).
Sister palaeodemes— 1 only.
Hla 213 Die elo: 3 intact cones.
Specific diagnosis
A Rissikisftvbus species with an ovuliferous scale consisting of 3 nearly
similar-sized lobes.
Specific characters
Strobilus: long (to ca 80 mm).
Megasporophyll: bracts distinctly longer than scales; scales 3-lobed, with
outer lobes slightly reduced.
Ovule: tear-shaped {ca 1 X 0.5 mm).
Etymology
plenus (Lat.)— full, complete, with reference to the lobed scales.
Comment & comparison
The bract/scale complex in this species is particularly difficult to inter-
pret. The R4 reconstructions (tfs 4, 5, adjacent) are based on the two most
explicit specimens of the reference palaeodeme. Each shows different fea-
tures. In the holotype. PRE/F/6774a,b (tf. la,b, adjacent), the three ovule
pairs are seen clearly preserved in several places, as are the lanceolate
bracts. In PRE/F/6775 [tf. 2a,b, adjacent; pi. 20(6-8)], the distinctively
fluted, semicircular, distal halves of the three scale lobes are seen fortui-
tously preserved in only one area (a further specimen, PRE/F/6759b, pi.
20(3-5), shows the feature less clearly.) Apparently in the fully mature
specimens, or in fossilisation, the distal lamina of the scale detaches leaving
the three pairs of ovules— still attached to the spreading stalk— exposed.
It is possible that the two Umkomaas (Umk 111) species, R. plenus and
R. semireductus, represent parts of a morphological continuum or onto-
genetic series and should be combined as one taxon. There does exist a very
wide morphological range of Rissikia foliage (And. & And. 1989, pis
265-271 ) at both Umk 1 1 1 and Hla 213 (also including both Rissikistrobus
species) which may support the two-species option preferred here. Also,
the recognition of a third species, R. reductus , characterised by the lone
central lobe, does suggest the likelihood of a morphologically intermediate
species.
Rissikistrobus
FINALES
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107
Rissikistrobus semireductus J.M.And. & H.M.And., sp. nov.
Holotype
Specimen : PRE/F/22508; pi. 21(1-5).
Assemblage (TC): Umk 111 Die 2spp; Umkomaas Valley.
Preservation', incomplete cone, without counterpart, longitudinal outer
view showing several scales and a few in situ ovules; compression in
thinly laminated, carbonaceous (good cuticle) moderately baked, dark
grey shale with good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens'. 41 indivs (4 intact cones, 37 partial cones).
Sister palaeodemes— 2 only.
Hla 212(1 intact & 1 partial cone).
Hla 213 (2 intact cones).
Specific diagnosis
A Rissikistrobus species with an ovuliferous scale consisting of a large
central lobe and two very reduced lateral lobes.
Specific characters
Strobilus: long (to ca 85 mm).
Megasporophyll : bracts distinctly larger than scales; scales 3-lobed, with
outer lobes strongly reduced.
Ovules: linear-elliptical {ca 2 X 0.5 mm).
Etymology
semireductus (Lat.)— with reference to the semireduced lateral lobes.
Comments & comparison
The structure of the R. semireductus bract/scale complex in external
and lateral view is most clearly visible in the holotype. The two best pre-
served units, as indicated in tf. 1 adjacent, are reconstructed (R3) at higher
magnification (tfs 2, 3) in abaxial and adaxial view. The additional 40
cones and cone fragments included in this species do not add a great deal
and some may well belong to R. plenus.
Rissikistrobus reductus J.M.And. & H.M.And., sp. nov.
Holotype
Specimen : PRE/F/20648a,b; pi. 22(3-7).
Assemblage (TC): Aas 411 Dic/Sph; Aasvoelberg.
Preservation : fairly complete cone (proximal end missing), part and counter-
part, longitudinal outer view and part section showing many scales and
a few in situ ovules; 3D mould and cast, imperfectly preserved;
impression in thinly laminated, strongly baked, yellowish grey shale
with very good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 20 indivs (2 complete & 18 partial cones).
Sister palaeodemes— nil.
Specific diagnosis
A Rissikistrobus species with an ovuliferous scale consisting of a single
lobe.
Specific characters
Strobilus: relatively short (to ca 60 mm).
Megasporophyll: bracts reduced, far shorter than scales; scales unlobed
[(?)outer lobes lost].
Ovule: linear-elliptical {ca 0.8 X 0.35 mm).
Etymology
reductus (Lat.)— with reference to the completely reduced lateral lobes.
Comments & comparison
The type locality, Aas 411, has yielded a good palaeodeme of Rissiki-
strobus cones that are quite distinct from the Umk 111 specimens. We are
obviously dealing with a separate species, though this is not evident in the
affiliated Rissikia foliage.
PINALES
Rissikistrobus
10B
d/ TRELITZIA 15 (2003)
Rissikianthus J.M.And. & H.M.And., sp. nov.
Type species
Rissikianthus townrowii J.M.And. & H.M.And., sp. nov.
Generic diagnosis
A pinalean male cone of small size (ca 10-18 mm long) with micro-
sporophylls bearing a pair of horizontally aligned abaxial microsporangia.
Generic characters
Strobilus: a compact cone, lanceolate to elliptical, of small size (10-18 mm
long); axis relatively gracile, strongly flexed towards base, with short
to long foliated free end; microsporophylls helically attached, in 7-10
gyres of ca 6-12 units (at midlength).
Microsporophyli. a simple scale, proportionately intermediate in length,
more or less straight, at 90° from axis; distal lamina woody, variously
triangular, entire, with strong heel; stalk broadly tapering proximally,
winged and keeled; microsporangia abaxial, two in number, extending
horizontally from heel of distal lamina, sessile.
Microsporangium: rotund (ca 2 X 1 mm), shallowly longitudinally
bilobed.
Pollen: striate disaccate.
Etymology
Rissikianthus— emphasising affiliation with the foliage Rissikia.
Global range: 4 spp., Gondwana, Tr. (CRN).
First & last: the 4 Molteno species described here.
Gondwana Triassic occurrence
SAf— Karoo Basin, 5 TCs (79 indivs).
Molteno occurrence
Frequency (F): 5 TCs (of 100 sampled in the Molteno).
Diversity (D): 4 species.
Abundance (A): 79 indivs total, rare to extremely rare.
Pen 321 Dic/Ris: 30 indivs
Kap 111 Dic/Ris: 25
Boe 112 Die cor: 1
Aas 411 Dic/Sph: 15
Umk 111 Die 2spp: 8
While the three genera of Molteno voltzialean male cones are all
extremely rare, those of Rissikianthus are markedly more frequent and
common.
Affiliated organs
Female strobilus: Rissikistrobus— Grade 4 (Kin. reinf., Mut. occ.).
Foliage: Rissikia— Grade 4 (Kin. reinf., Mut. occ.).
Though nowhere known in organic connection, the affiliation of
Rissikistrobus, Rissikianthus and Rissikia is considered virtually certain
(see further on pp. 104, 105).
Classification & comparison
Suprageneric classification (Podocarpaceae/Pinales)
Townrow (1967) records that cones with two pollen sacs and disaccate
pollen as in Rissikianthus (his Rissikia) are found with regularity only in
the Podocarpaceae and Pinaceae. With regard to the pollen, he records that
‘the corpus and saccus shape and ornamentation seem .... to come close to
Dacrydium.' He states further that striae, which characterise the
Rissikianthus pollen ‘are not known in any living conifer.’ Townrow con-
cludes, along with evidence from the female cone and the foliage, that the
Rissikia/Rissikistrobus/Rissikianthus plant represents the earliest known
member of the Podocarpaceae (see fuller discussion on p. 105).
Intergeneric comparison (Molteno genera)
Five pinopsid (coniferopsid) male cone genera have been recognised
here in the Molteno: Fredianthus, Lutanthus and Odyssianthus included in
the Voltziales; Rissikianthus in the Pinales; and Helvetianthus in an unde-
fined order. Rissikianthus is distinctive amongst these genera in being the
only form with typical pinalean scales bearing a pair of abaxial, horizon-
tally aligned microsporangia. The three voltzialean genera all bear
microsporangia attached to the scale stalk, while Helvetianthus is a very
different strobilus without conventional scales or microsporangia.
n 35 man-hrs cleaving (10 per 1 man-day) rare
65 “ “ ( 3 “ 1 “ ) very rare
6 “ “ ( 1 “ 1 “ )
512 “ “ ( 1 “ 3 “ )
400 “ “ ( 1 “ 5 “ ) extr. rare
Reconstructions
In the reconstructions of the four Rissikianthus species we attempt a
faithful portrayal of the norm for each of the reference palaeodemes (from
Pen 321, Aas 411, Umk 1 1 1 and Kap 111). Definite limitations have to be
emphasised, however.
R. concavus (R3-4)— The Pen 321 palaeodeme shows excellent 3D
(mould/cast) preservation in many specimens, such that the reconstruction
is particularly reliable. The size and shape of the cones is remarkably con-
stant, falling within a particularly narrow range of variation. This might be
explained by their having occurred within a very limited area (<1 m along
strike) of the thin chert bed in which they were found. They may even rep-
resent a single tree and a single season’s growth. The microsporangia are in
several instances clearly distinguished— in being notably delicate with a
transparent bluish tinge in contrast to the more rusty brown woodiness of
the cone scales.
R. linearis , R. townrowii, R. convectus (all R4) — The reference palaeo-
demes of these three species are far less clearly preserved. The outline
shape of the scales, one of the diagnostic features in recognising species,
are nowhere unambiguously seen in the available specimens.
Rissikianthus
PINALES
TRELITZIA 15 (2003)
109
Intactness & preservation of cones (Molteno)
The small Rissikianthus cones— as for their larger Rissikistrobus ovu-
late counterparts — are all preserved essentially intact. This holds for each
of the five TCs in which they occur.
Microsporangia
It would appear that the normal condition of preservation of the
Molteno Rissikianthus cones, in addition to being intact, is with their sac-
like microsporangia in place. Each of the palaeodemes is briefly discussed:
Pen 321 Dic/Ris (30 indivs)— The microsporangia are best seen in the
Pen 321 palaeodeme which has yielded a good number of mature cones
from within a single closely confined pocket in the extensive cherty-mud-
stone horizon of Peninsula. These cones are exquisitely preserved as
moulds/casts, mostly in longitudinal view, partly in section showing the
delicate microsporangial sacs, partly in outer perspective showing clearly
the shape of the scale face. Two cones [pi. 24(3, 4)] are preserved perfect-
ly in cross-section displaying the microsporangia in plan view. Although all
specimens from Pen 321 probably bear in situ microsporangia, they are
particularly well seen in only seven of the 30 or so cones.
Aas 411 Dic/Sph (15 indivs)— Here the microsporangia are preserved
as moulds/casts, much as at Pen 321, but occur in thinly laminated, strong-
ly baked shale and are not as clearly seen. Most specimens show the axis
and scales in longitudinal section and, in five of the 15 cones, in situ
microsporangia are more or less clearly evident.
Umk 111 Die 2 spp (8 indivs)— In contrast to Pen 321, and particular-
ly Aas 411, the Rissikianthus cones from Umk 111 are preserved (as com-
pressions in carbonaceous shale with potential cuticle) in outer view with
the scales fully closed and overlapping. In two instances, the cones are seen
largely in section showing (none too clearly) in situ microsporangia. As
shown by Townrow (1967), the compression material from Umk 1 1 1 does
yield cuticular material and in situ pollen.
Kap 111 Dic/Ris (25 indivs)— This relatively numerate palaeodeme
derives from a thinly laminated, medium dark grey shale. The cones are
seen partly in section, partly in outer view, but are not clearly preserved.
The full pollen sacs are evident in some five instances.
Roe 112 Die cor (1 indiv)— The single specimen from this TC, though
imperfectly preserved, shows in situ pollen sacs.
Cuticles (& in situ pollen)
Potential sample: Umk 111,8 indivs.
Macerated (this work): nil.
Preser\’ation grade: Grade 4—5.
Diagnostic characters: see Townrow (1967); this vol., p. 105.
In situ pollen: striate disaccate grains.
Comment: The grade and characters outlined above are based exclusively
on Townrow (1967). We have not further studied the cuticle of the cones.
Significance:
Classification— The cuticle of Rissikianthus (and its affiliated organs)
does not contribute unambiguously, in our knowledge, to classification
within the Pinopsida in general or the Pinales in particular (but see
Townrow’s reflections quoted on p. 105).
Affiliations— Townrow (1967) established to his satisfaction that the
cuticle of the affiliated organs (foliage and cones) corresponded. Some of
his drawings are refigured on p. 105. For foliage cuticles, see p. 112.
Adaptive radiation (Molteno diversity)
Assessment of diversity within Rissikianthus is particularly dependent
on the clarity of preservation of the cones. Interpretation of critical diag-
nostic morphology is not always certain. Acknowledging the limitations,
we find each of the four available palaeodemes (the single specimen from
Boe 112 is not considered) to show nonoverlapping morphological varia-
tion and, therefore, to represent four separate species. It is in the shape of
the scales (distal lamina), the number of scales per gyre and overall shape
and size of the cones that the species are characterised.
Should the diversity —of four species— be real, it is not reflected in the
foliage where only a single variable species can be recognised in the four
relevant TCs. Comparison of diversity with the affiliated female cones
(Rissikistrobus) is complicated by the fact that only two well-represented,
reasonably preserved palaeodemes (from Umk 111 and Aas 411) are at
hand (Tab. 38). The collections from each of these sites, however, do indi-
cate two distinctive species for both the male and female cones.
The four Molteno species are based on the TCs/reference palaeodemes
as indicated below. Each derives from a distinct habitat and from a differ-
ent level within the stratigraphic sequence— which lends support to the
recognition of the four taxa.
R. concavus— Pen 321 Dic/Ris (Peninsula), 30 indivs
Dicroidium open woodland; Cycle 2f (Indwe Member)
R. linearis— Aas 411 Dic/Sph (Aasvoelberg), 20 indivs
Sphenobaiera closed woodland; Cycle 1 (Bamboesberg Member)
R. townrowii—Xlrak 111 Die 2spp (Umkomaas Valley), 8 indivs
Dicroidium riparian forest (mature); Cycle 2b (Indwe Member)
R. convectus— Kap 111 Dic/Ris (Kapokkraal), 25 indivs
Dicroidium riparian forest (immature); Cycle 2e (Indwe Member)
GONDWANA TRIASSIC, GEOSTRAT DISTRIBUTION
PINALES
Rissikianthus
110
d^TRELITZIA 15 (2003)
Rissikianthus concavus J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/13674 a,b‘x’; pis 23(2, 7, 8), 24(2).
Assemblage (TC): Pen 321 Dic/Ris; Peninsula.
Preservation: complete cone (missing proximal half of free axis), part and
counterpart, longitudinal outer view and section with in situ micro-
sporophylls; ca 60% flattened, exquisitely preserved; 3D mould and
cast in thickly laminated, medium light grey shale with poor cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 30 indivs (intact strobili, partial strobili), pis 23( 1-10), 24(1M),
25(1-7).
Sister palaeodemes— nil.
Specific diagnosis
A Rissikianthus cone of small size and narrowly elliptic shape bearing
ca 7 sharply concave-sided scales per gyre, and borne on a particularly long
free gracile axis.
Specific characters
Strobilus: small (ca 10 X 3.4 mm), narrowly elliptical, apex acute, axis gracile,
free axis particularly long (to 24 mm), scales ca 7 per gyre.
Microsporophyll: distal lamina with deep depression at insertion of stalk,
commissure (line of junction) a very pronounced groove.
Etymology
concavus (Lat.) — with reference to the concave scale margin.
Comment & comparison
Of the four species of Rissikianthus recognised here, R. concavus is the
most common and by far the most clearly preserved and distinctive. The
reference palaeodeme of 30 individuals shows the cone exquisitely in outer
view, longitudinal section and cross section.
Rissikianthus linearis J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/20644 a.b; pi. 26(1-3).
Assemblage (TC): Aas411 Dic/Sph; Aasvoelberg.
Preserx’ation: complete cone (missing proximal end of free axis), part and
counterpart, longitudinal section, without microsporangia; impression
in thinly laminated, strongly baked, yellowish grey shale with very
good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 15 indivs (intact strobili, partial strobili), pi. 26(1-9).
Sister palaeodemes— nil.
Specific diagnosis
A Rissikianthus cone of relatively large size and oblong elliptic shape
bearing ca 6 weakly concave-sided scales per gyre, and borne on a short
gracile free axis.
Specific characters
Strobilus: relatively large (ca 14 X 4.4 mm), oblong elliptical, apex obtuse,
axis gracile, free axis medium, scales ca 6 per gyre.
Microsporophyll: distal lamina glabrous, outer margins straight to weakly
convex, with moderate depression at insertion of stalk, commissure
(line of junction) a clear linear groove.
Etymology
linearis (Lat.)— with reference to the linear shape of the strobilus.
Comment & comparison
This species compares most closely with R. concavus in various diag-
nostic features and differs from it most evidently in size and length of free
axis. It is interesting that the four Rissikianthus species each represent a dif-
ferent habitat within the Molteno plain (Tab. 38) and that R. linearis and R.
concavus occurred in woodland rather than forest.
Rissikianthus
PINALES
c/trELITZIA 15 (2003)
Rissikianthus tOWWOWii J.M.And. & H.M.And., sp. nov.
Holotype
Specimen : PRE/F/6765a; pi. 27(7, 8).
Assemblage (TC): Umk 1 1 1 Die 2spp, Umkomaas Valley.
Preservation', complete cone, primarily in longitudinal section, with micro-
sporangia in situ; compression in thinly laminated, carbonaceous (good
cuticle) moderately baked, dark grey shale with good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens'. 8 indivs (mostly complete intact strobili), compressions with
cuticle; pi. 27(1-8).
Sister palaeodemes— nil.
Specific diagnosis
A Rissikianthus cone of relatively large size and roundly elliptic shape
bearing ca 12 finely striate, straight-sided scales per gyre, and borne on a
short stout free axis.
Specific characters
Strobilus: relatively large (ca 14 X 7.4 mm), roundly elliptical, apex obtuse,
axis robust, free axis short, scales ca 12 per gyre.
Microsporophyll: distal lamina finely striate throughout apical part, outer
margins more or less straight, with shallow or no depression at inser-
tion of stalk, commissure (line of junction) not pronounced.
Eponymy
townrowii— in honour of Dr John Townrow, who (in 1967) first described
but did not name these male cones from the famous Umkomaas local-
ity.
Comment & comparison
R. townrowii and R. convectus are similar in the nature and number of
scales, but quite dissimilar to the other two Rissikianthus species. They
occur within the mature and immature Dicroidium riparian forest habitats
respectively (Tab. 38). R. townrowii. however, is distinct from R. convec-
tus in the roundly elliptic shape of the strobilus and ornamentation of the
exposed distal laminae of the scales.
Rissikianthus convectus J.M.And. & H.M.And., sp. nov.
Holotype
Specimen'. PRE/F/16227; pi. 28(4, 5).
Assemblage (TC): Kap 1 1 1 Dic/Ris, Kapokkraal.
Preservation: complete cone (missing free axis), without counterpart, lon-
gitudinal section, without microsporangia; impression in thinly lami-
nated, moderately baked, medium dark grey shale with very good
cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 25 indivs (intact strobili, partial strobili), pi. 28(1-7).
Sister palaeodemes— nil.
Specific diagnosis
A Rissikianthus cone of relatively large size and narrowly elliptic
shape bearing ca 10 partially striate, convex-sided scales per gyre, and
borne on a short moderately stout free axis.
Specific characters
Strobilus: relatively large (ca 17 X 4.4 mm), narrowly elliptical, apex obtuse,
axis moderately robust, free axis short, scales ca 10 per gyre.
Microsporophyll: distal lamina finely striate down middle, outer margins
convex, with shallow or no depression at insertion of stalk, commis-
sure (line of junction) not pronounced.
Etymology
convectus (Lat. ) — with reference to the convex scale margin.
Comment & comparison
R. convectus (from Kap 1 1 1 ), as noted above, is most like R. townrowii
(Umk 111), the other species occurring in Dicroidium riparian forest, and
is markedly distinct from the two woodland species. It differs from the
Umk 1 1 1 species most notably in the linearly elliptical shape of the stro-
bilus and the more restricted striated ornamentation on the scale face.
FINALES
Rissikianthus
112
d^TRELITZIA 15 (2003)
Rissikid Townrow 1967
Type species
Rissikia media (Ten. -Woods 1883) Townrow 1967.
Queensland, Australia; U. Triassic.
Generic diagnosis
A pinalean short shoot with linear single-veined leaflets that are heli-
cally attached and twisted near their base giving an overall appearance of a
planar pinnate leaf.
Generic characters
Attachment, unknown.
Foliage shoot: short shoot individually dehisced, small, finely linear to
elliptical; leaflets flattened, linear to narrowly elliptical, helically
attached but twisted near base to extend in one plane on either side of
rachis, adpressed to spreading at 80° from rachis, apex obtusely to
acutely pointed, base clearly contracted or not contracted; with a single
median vein.
Cuticle (adapted from And. & And. 1989); this vol., tfs 1^4 below.
Yield: Lit 111—15 specimens sampled, grade index 15/0/0/0/0.
Umk 111 — 19 specimens sampled, grade index 10/3/6/0/0.
Diagnostic characters (based on R. media, Umk 111)
Adaxial/abaxial: isobilateral, U & L cuticle of equal thickness.
Cell characteristics: narrowly oblong, end walls oblique to square; U & L
cuticle with 450 cells per mm2; walls straight to gently curved, profile
triangular, plan gracile; cells over veins nondistinctive, noncutinised;
occasionally faintly papillate.
Stomatal apparatus: haplocheilic, amphistomatic, U & L cuticle with ca
20 stomata per mm2, nonaligned, longitudinal; subsidiary cells
(anomo) brachyparacytic, 2 or 3 in number, noncutinised, radial walls
normal; guard cells narrowly elliptic, 36 X 14 mp, polar sulcus deep,
non-omamented, labia gracile; Florin ring lappetate, lappets flattened,
arching; stomatal pit unknown.
Other features: none preserved.
Eponymy
Rissikia— after the family Rissik who, in the 1950s & 1960s, owned the
land neighbouring that on which the Umkomaas locality (Umk 111) is
sited.
Global range: 2 species, Gondwana, Tr. (ANI-NOR).
First: Rissikia media (Holmes 1982); Benolong, Dubbo, Australia.
Last: Rissikia ( Elatocladus raoi ) (Pal 1984); Tiki Fm., S. Rewa/Tiki, India.
Gondwana Triassic occurrence
Frequency (F): 17 degree squares (of the 84 across Gondwana).
Ubiquity (U): 5 continents (of 5 comprising Gondwana).
Diversity (D): 2 foliage species (as recognised in And. & And. 1989).
Abundance (A): 1% (the norm as in Molteno TCs).
Longevity (L): 14 myrs (late Anisian to early Norian).
Colonisation success: FUDAL rating 17/5/2/1/14 = 39.
Intermediate success (Grade 3): Rissikia was the 1 1th most prominent
genus in the Gondwana Triassic; it was frequent, ubiquitous and fairly
long-lived, but low in abundance and markedly lacking in diversity.
Endemism: of the two described Gondwana Triassic species, the rarer
(R. eskensis) has a disjunct distribution (Karoo Basin, and Clarence-
Moreton Basin, Australia), while the second (R. media) occurs scat-
tered throughout the realm.
Molteno occurrence
Frequency (F): 21 TCs (of 100 sampled in the Molteno).
Diversity (D): 2 species.
Abundance (A); co-dominant (35-38%) in 2 TCs, occasional (1-2%) in 5
TCs, and <1% in the other 14 TCs.
Habit: probably a large tree.
Preferred habitat: Dicroidium riparian forest or Dicroidium open wood-
land, occasionally forming monodominant wetland stands.
Affiliated organs
Female strobilus: Rissikistrobus — Grade 4 (Kin. reinf., Mut. occ.).
Male strobilus: Rissikianthus— Grade 4 (Kin. reinf., Mut. occ.).
Classification & comparison (adapted from And. & And. 1989, p. 450)
Intergeneric comparisons
Gondwana Triassic pinopsid genera— The foliage of Rissikia, with its
helically attached linear leaflets twisted near their base to spread in one
plane, is quite distinct from that of the other four pinopsid genera found in
Gondwana Triassic strata. Voltziopsis and Pagiophyllum leaflets are small,
scale-like, distinctly thickened and unveined; while those of Heidiphyllum
and Clariphyllum are relatively large, flattened and multiveined.
Other pinopsid genera— Elactocladus Halle (1913) is a global Meso-
zoic form-genus which is much like Rissikia. It differs principally in that
its leaflets are spirally borne and strongly contracted at the base to form a
short petiole attaching the leaf to a basal cushion.
Interspecific comparisons
The two species of Rissikia recognised here from the Gondwana
Triassic fall, on the basis of leaf macromorphology, readily within the com-
pass of a natural genus. However, affiliated cones (female and male) and
cuticle are known only for R. media, so it is not possible to confirm the true
generic relationship between the two species.
The reference palaeodemes of the two species are perfectly distinctive.
A good number of additional palaeodemes, particularly from Australia,
exist of both species and from the data at hand their appearance in the var-
ious assemblages appears to be virtually mutually exclusive. Resolving
whether this is really true for the Australian material and whether the
species are always distinct, must await more comprehensively published
data.
R. media
2
Rissikia
PINALES
c/tRELITZIA 15 (2003)
113
Rissikia
l!mk 111
tfs 1-9
from And. & And., 1989
Kon 222
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
PINALES
Rissikia
114
(2003)
Umkomaas Valley
(Umk 1 1 1 Die 2spp)
all PRE/F/6774a,b
Holotype
Rissikistrobus plenus
pi. 19
PINALES
115
^ TRELITZIA 15 (2003)
& 1 i
BP/2/1148
BP/2/1148
PRE/F/6775 £*
Umkomaas Valley
(Umk 111 Die 2spp)
PRE/F/6759a
PRE/F/6759a
PRE/F/6759b
PRE/F/6759a
PRE/F/6759a
PINALES
pi. 20
Rissikistrobus plenus
116
g/trelitzia 15 (2003)
PRE/F/785
PRE/F/22508
Umkomaas Valley
(Umk 1 1 1 Die 2spp)
PRE/F/22508
Holotype
Holotype
pi. 21
Rissikistrobus semireductus
PINALES
RELITZIA 15 (2003)
Holotype
PRE/F/20849a
Aasvoelberg
(Aas 411 Dic/Sph)
PRE/F/20849a
PRE/F/20648b
PINALES
pi. 22
Rissikistrobus reductus
118
r/TRELITZIA 15 (2003)
PRE/F/13674a
Holotype
Peninsula
(Pen 321 Dic/Ris)
PRE/F/13672b
PRE/F/13672a
PRE/F/13680
PRE/F/13672a
PRE/F/13674a
PRE/F/13674a'x'
X10
PRE/F/13675b
X10
Rissikianthus concavus
pi. 23
PINALES
^TRELITZIA 15 (2003)
Peninsula
(Pen 321 Dic/Ris)
PRE/F/13740b
PRE/F/13674'x' X20 i
PRE/F/13674a'y'
FINALES
pi. 24
Rissikianthus concavus
120
d^TRELITZIA 15 (2003)
PRE/F/13680
PRE/F/22153
Rissikianthus concavus
pi. 25
PINALES
Peninsula
(Pen 321 Dic/Ris)
X20
121
r
RELITZIA 15 (2003)
Holotype
Aasvoelberg
(Aas 411 Dic/Sph)
PRE/F/21742b
PINALES
pi. 26
Rissikianthus linearis
122
d?TRELITZIA 15 (2003)
PRE/F/6765a
Holotype
Umkomaas Valley
(Umk 1 1 1 Die 2spp)
PRE/F/6408b
PRE/F/6408a
Rissikianthus townrowii
pi. 27
PINALES
r^f- TRELITZIA 15 (2003)
PRE/F/16228
PRE/F/16228
Kapokkraal
(Kap 1 1 1 Dic/Ris)
PINALES
pi. 28
Rissikianthus convectus
124
^VtreLITZIA 15 (2003)
PINOPSIDA S.V.Meyen 1984
PIN ALES S.V.Meyen 1984
INCERTAE SEDIS family
Pagiophylllim Heer 1881
l^pe species
Pagiophylllim circinicum (Saporta) Heer 1881.
Sierra de San Luiz. Portugal; U. Triassic (Malm).
Generic diagnosis: see Hams (1979).
Generic characters (after Harris 1979)
Foliage shoot: 'Shoot, bearing leaves in a helix; leaf about as broad as its
basal cushion, length exceeding width of cushion. Blade not or scarce-
ly narrowed at its base, in section broader horizontally than vertically’.
Cuticle: see And. & And. (1989, p. 465) for P. bosei (India) cuticle; this
vol., tf. 3 below.
Etymology
Pagiophylllim— pagio (Gr.), firm, solid; phyllum (Gr.), leaf.
Global range: numerous spp.. Pangaea-wide, Palaeozoic to recent.
Gondwana Triassic occurrence
Frequency (F): 4 degree squares (of the 84 across Gondwana).
Ubiquity (U): 4 continents (of 5 comprising Gondwana).
Diversity (D): 4 foliage species.
Abundance (A): <1% (the norm in Molteno TCs).
Longevity (L): 6 myrs.
Colonisation success: FUDAL rating 4/4/4/— /6 = 18.
Intermediate success (Grade 3): Pagiophylllim was the 12th most
prominent genus in the Gondwana Triassic; it was of fairly low fre-
quency. ubiquity, diversity, abundance and longevity. (Note that in Tab.
20 (p. 27), the hierarchical position is given as 21 and the FUDAL rat-
ing as 12. The reason for this is that the South American and Trans-
Antarctic records were not yet included in the table.)
Endemism: all species are single-assemblage endemics.
Molteno occurrence
Frequency (F): 1 TC (of 100 sampled in the Molteno).
Diversity (D): 1 species.
Abundance (A); extremely rare at a single TC.
Habit: probably a tree.
Preferred habitat: possibly preferring Sphenobaiera open woodland.
Affdiated organs
Female strobilus: unknown.
Male strobilus: unknown.
Classification & comparison (adapted from And. & And. 1989, p. 464)
Intergeneric comparisons
Gondwana Triassic pinopsid genera— Pagiophylllim is alone among
the five pinopsid foliage genera (aside from Clariphyllum) found in the
Gondwana Triassic in that it is a form-genus for which the cones remain
unknown. The foliage shoots and leaflets are in many ways like those of
Voltziopsis, but the cuticle of the latter is distinctly different (from the
species P. bosei) in that its subsidiary cells are anomocytic to actinocytic,
noncutinised and nonlappetate.
Other pinopsid genera— Brachy phyllum (Lindley & Hutton 1836) is a
form-genus much confused with Pagiophylllim. It differs (after the diag-
noses of Harris 1969, 1979) principally in that the free apical part of the
leaflet is less than the width of the basal cushion, while in the latter it
exceeds the width of the cushion.
Interspecific comparisons
As noted adjacent, Pagiophylllim is a pandemic form-genus incorpo-
rating numerous species from the Palaeozoic to recent. The foliage type is
found associated with entirely different cones falling in widely separated
conifer families such as the Cheirolepidiaceae and Taxodiaceae.
The two Gondwana Triassic species, including that now described
from the Molteno, are extremely rare, being known only from their sparse
reference palaeodemes— four illustrated individuals for P. bosei (India) and
two individuals for P. aasvoelense (Molteno). [For more recent data on
South American material, see Spalletti et al. ( 1991 ), and on Antarctic mate-
rial, see Cantrill et al. (1995).]
Cuticle
The Pagiophylllim cuticle (based on the Indian species P. bosei) shows
very different features from that of the other Molteno pinopsid genera. It
has a thick cuticle with subsidiary cells that are regularly actinocytic,
cutinised and strongly lappetate. Dordrechtites , Heidiphyllum and Rissikia
(plus its reproductive affiliates) have thin cuticles with anomocytic or
brachyparacytic subsidiary cells that are mostly nonlappetate. This would
suggest that Pagiophylllim most probably falls in a position within pinop-
sid classification quite remote from the other Molteno genera.
Pagiophylllim
PINALES
(§f‘ TREL1TZI A 15 (2003)
125
Pagiophyllum aasvoelense j.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/20820; pi. 29(3), tf. 1 adjacent.
Assemblage (TC): Aas411 Dic/Sph; Aasvoelberg.
Preserx’ation: impression in thinly laminated, strongly baked, yellowish
grey shale with very good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 2 individuals.
Sister palaeodemes— nil.
Specific diagnosis
A Pagiophyllum species with small ( ca 2 X 0.1 mm) rhomboid to tri-
angular leaflets.
Specific characters
Stem: ca 0.44 mm wide.
Leaf blade: rhomboid to triangular in shape, size varies from leaves 2 mm
long in holotype to much smaller, 1 mm long, in the second individual.
The attachment appears to be helical, with the leaves spread sideways.
Etymology
aasvoelense— after the type locality.
Comment & comparison
This species is similar to P. bosei (tfs 1-3 opposite) from India but the
leaves are not strongly keeled. P. papillatus from Antarctica has leaves
clearly helical, lanceolate and much larger ( 1-5 X 1-3 mm).
PINALES
Pagiophyllum
126
d^TRELITZIA 15 (2003)
PINOPSIDA S.V.Meyen 1984
INCERTAE SEDIS order
INCERTAE SEDIS family
Gypsistrobus J.M.And. & H.M.And., gen. nov.
Type species
Gypsistrobus scutatus J.M.And. & H.M.And., sp. nov.
Generic diagnosis
A pinopsid female cone of linear-elliptic shape with a gracile axis and
single-lobed bract/scale complexes bearing single bilobed ovules.
Generic characters
Strobilus: cone compact, linear, small (to >35 X 6 mm); axis gracile (ca
0.8 mm diam.). free end ca 5-12 mm long, gently curving; megas-
porophylls spirally arranged, ca 6-8 units per gyre.
Megasporophyll: cone units consisting of ovuliferous bract/scale complex-
es; bracts free, leafy, lanceolate, ca 5 mm long; scales ovate, acute-
tipped; ovules/seeds adaxial, single, bilobed, in concave surface of
scale.
Ovule/seed: elongately oval, bilobed, 1-2 mm long.
Etymology
Gypsistrobus— gyps (Gr.), vulture, with reference to the type locality Aasvoel-
berg, which means ‘vulture mountain’ in Afrikaans; strobilos (Gr.), cone.
Global range: 1 sp., Gondwana, Tr. (CRN).
First & last: the single Molteno species described here.
Gondwana Triassic occurrence
SAf— Karoo Basin, 1 TC (5 indivs).
Molteno occurrence
Frequency (F): 1 TC (of 100 sampled in the Molteno).
Diversity (D): 1 species.
Abundance (A): 5 individuals.
Aas 41 1 Dic/Sph: 5 indivs in 512 man-hrs cleaving (1 per 10 man-days), extr. rare
Affiliated organs
Female strobilus— unknown.
Foliage— unknown.
Within the pinopsid foliage and male-cone material preserved in the
Aas 411 TC, there remain Pagiophyllum, Fredianthus and Lutanthus (L.
robustus) without female-cone affiliates. These three genera appear to be
most readily placed in the Pinales and Voltziales and are less likely to link
with Gypsistrobus.
Classification & comparison
Suprageneric classification (family & order Incertae)
As for the following genus, Avistrobus (p. 130), we have made no
attempt to include Gypsistrobus in our ‘Global classification of the gym-
nosperms’ (And. & And., in prep.). They most likely represent a single
undescribed family and order of pinopsid.
The closest comparison would appear to be with Ferugliocladus
(Ferugliocladaceae) of Archangelsky & Cuneo (1987) from the Permian
(Asselian) Rio Genoa Group, Central Patagonian Basin, Argentina. This
Permian genus bears megasporophylls with a large, triangular, free bract
and single, sessile, orthotropous, fully enclosed ovules. We interpret the
two Molteno genera as having naked ovules, but the material is insuffi-
ciently preserved to verify this.
Further somewhat similar ovulate genera are Krylovia and Suchoviella
(order Cordaitanthales, family Rufloriaceae) from the Carboniferous-
Permian of the former USSR. These genera both bear naked uni-ovulate
scales superficially like Gypsistrobus, but with definite micropyles and no
free bracts.
Intergeneric comparison (Gondwana Triassic)
Gypsistrobus is similar in size and shape of strobilus to Rissikistrobus
re ductus but differs in several respects: the ovules appear to be bifidly
lobed rather than occurring in pairs; the megasporophylls dehisce readily
and they show no evidence of persistent reduced foliage on the petiole.
Avistrobus, known only from a single specimen from Birds River (Bir
111), appears comparable to Gypsistrobus at family level. It differs most
evidently in the scales bearing single ovules with no evidence of being lon-
gitudinally bilobed, and in the far more robust nature of the petiole and
axis.
Reconstructions
The reconstructions are based on the holotype and two sister speci-
mens of the reference palaeodeme (tfs 1-3 opposite).
In view of the imperfect preservation, the exact morphology of the
bract/scale complex cannot be certainly established. Indeed, it is only in
PRE/F/21736 (tfs 1, 2 above), interpreted as an immature cone, that the
leafy bracts are seen. They appear to have been entirely lost in the four
mature cones where the ovuliferous scales are seen in various states of dis-
articulation.
The cones are all shown in pendent orientation. The slender, seeming-
ly herbaceous axes, could hardly have held these cones erect.
Gypsistrobus
PINOPSIDA/INCERTAE
dS^TRELITZI A 15 (2003)
127
Gypsistrobus SCUtatUS J.M.And. & H.M.And., sp. nov.
Holotype
Specimen-. PRE/F/21090a,b; pis 30(3, 4, 8, 9), 31(1).
Assemblage (TC): Aas411 Dic/Sph; Aasvoelberg.
Preservation: fairly complete cone (apex and proximal end of axis missing),
part and counterpart, longitudinal outer view with many scales;
impression, imperfectly preserved; impression in thinly laminated,
strongly baked, yellowish grey shale with very good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 5 indivs (4 intact; 1 partial).
Sister palaeodemes— nil.
Specific diagnosis— as for genus.
Specific characters— as for genus.
Etymology
scutatus— scutum (Lat.), shield, with reference to the shape of the scale.
Classification & comparison — see notes for genus.
Intactness & preservation of cones
The five G. scutatis cones comprising the reference palaeodeme appear
to represent a series showing progressive disarticulation.
PRE/F/21736a,b (tf. 1)— This we interpret to be an immature cone
with fully intact scales along the distal half of the axis, with relatively small
scales and only partially developed seeds, and with the bracts variously
tom and shredded yet still in place. Were it not for this cone, we would have
no knowledge of bracts characterising this taxon.
PRE/F 21090a, b (tf. 2)— The scales in this second cone are largely
detached, yet still more or less in place, while the seeds are distinctly fuller
and apparently mature.
PRE/E21091a,b (tf. 3)— This third figured specimen also appears
mature, with full seeds and scales that presumably disarticulated to their
relatively dispersed state after deposition.
PRE/F/13099 & PRE/F/21328 (not figured as pen sketches)— The
fourth and fifth specimens in the collection show similar or still further levels
of disarticulation. PRE/F/13099 (not illustrated) is a more or less complete
cone and set of scales in a state of post-deposition disarticulation much like
that shown in tf. 2, while PRE/F/21328, pis 30(5, 6), 31(5, 6), is a cluster
of scales dissociated from their parent cone.
Molteno occurrence (elaborated)
For a full account of the Aas 411 TC. in which Gypsistrobus is unique,
see And. & And. (in prep.) (a sequal to this work).
Habit & habitat
Given the rarity of Gypsistrobus, it is most likely an allochthonous
element within the Aas 411 TC (a lake deposit in the floodplain). In the
absence of foliage or male-cone affiliates, it is likely to represent a rela-
tively rare element within the floodplain community.
The Aas 41 1 TC occurs low in the Molteno Fm. (Cycle 1) and appears
to include a few possibly more primitive pinopsid taxa (e.g. Fredianthus
and Lutanthus robustus) than the younger horizons.
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
INCERTAE/PINOPSIDA
Gypsistrobus
128
rV TRELITZIA 15 (2003)
PRE/F/21328
'mmm
PRE/F/21091a /'/ X2
X2
PRE/F/21736b
jfj PRE/F/21090a ||
Holoytpe
PRE/F/21090b
Aasvoelberg
(Aas 41 1 Dic/Sph)
PRE/F/21090a
PRE/F/21090b
PRE/F/21091a
Gypsistrobus scutatus
pi. 30
PINOPSIDA/INCERTAE
129
r/TRELlTZIA 15 (2003)
PRE/F/2 1736a
V PRE/F/21091a
Aasvoelberg
(Aas 411 Dic/Sph)
PRE/F/21736b
tit- f'' ' %
• PRE/F/2 1090a ^
frrrirBiM'rcr* bb
PRE/F/21328
PRE/F/21328
Gypsistrobus scutatus
INCERTAE/PINOPSIDA
130
^TRELITZIA 15 (2003)
PINOPSIDA S.V.Meyen 1984
INCERTAE SEDIS order
INCERTAE SEDIS family
AvistrobllS J.M.And. & H.M.And., gen. nov.
Type species
Avistrobus foliosus J.M.And. & H.M.And., sp. nov.
Generic diagnosis
A pinopsid female cone of elliptic shape with a stout axis and single-
lobed bract/scale complexes bearing single unlobed ovules.
Generic characters
Strobilus : cone compact, oval, small (to >25 X 10 mm); axis robust (ca 1.5
mm diam.), gently curving; megasporophylls spirally arranged, ca 6-8
units per gyre.
Megasporophylls : cone units consisting of ovuliferous bract/scale com-
plexes; bracts free, leafy, elliptical, ca 5 mm long; scales elliptical,
acute tipped; ovules/seeds adaxial. single, in concave surface of scale.
Ovules: elliptical, with median rib, ca 1 mm long.
Etymology
Avistrobus— avis (Lat.), bird, with reference to the type locality Birds River;
strobilos (Gr.), cone.
Global range: 1 sp., Gondwana, Tr. (CRN).
First & last: the single Molteno species described here.
Gondwana Triassic occurrence
SAf— Karoo Basin, 1 TC ( 1 indiv.).
Molteno occurrence
Frequency (F): 1 TC (1 indiv.).
Diversity (D): 1 species.
Abundance (A): 1 indiv.
Bir 1 1 1 2spp: 1 indiV. in 550 man-hrs cleaving ( 1 per 55 man-days), vanishingly rare
Affiliated organs
Female strobilus — unknown.
Foliage— unknown.
Classification & comparison
Suprageneric classification (family & order incertae)
For text relevant to both this and the previous genus, Gvpsistrobus, see
p. 126.
Intergeneric comparison (Gondwana Triassic)
Avistrobus , as noted previously (p. 126), has obvious similarities with
Gypsistrobus from Aas 411. If both have been interpreted correctly, then
they very likely represent the same family.
A second Molteno genus. Switzianthus (from Lit 111), is not dissimi-
lar in size and shape of strobilus, but is known to be male through having
yielded clear clusters of pollen grains. Cuticular comparison is excluded
since the Bir 1 1 1 floral assemblage consists purely of impressions on buff
shale.
Reconstruction
The reconstruction is based on the holotype (PRE/F/15591a, b), the only
specimen found to date representing the taxon. Sufficient morphological
detail is available to justify a sketch, though the bract/scale is certainly not
securely interpreted.
Molteno occurrence (elaborated)
For an analysis of the Bir 111 TC, to which Avistrobus is exclusive, see
And. & And. (in prep.), a sequel to the present work.
Habit & habitat
In view of its ‘vanishingly rare’ occurrence, Avistrobus is very likely
an allochthonous element within the Bir 111 TC (a lake deposit in the
floodplain). The plant was most probably a rare member of the floodplain
community. It is of interest that Avistrobus (Bir 111) and Gypsistrobus (Aas
411) occur high and low in the Molteno sequence, respectively, in the only
two thoroughly sampled lake-deposit TCs.
Avistrobus foliosus J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/15591a,b., pi. 32(1-6).
Assemblage (TC): Bir 111 Sph 2spp; Birds River.
Preservation: virtually complete cone (proximal end of free axis missing),
part and counterpart, longitudinal outer view with several scales seen
from different perspectives; impression in thinly laminated, yellowish
grey shale with very good cleavage.
Reference palaeodeme
Assemblage: as for holotype.
Specimens: the holotype.
Sister palaeodemes— nil.
Specific diagnosis— as for genus.
Specific characters— as for genus.
Etymology
foliosus (Lat.)— leafy, with reference to the papery leaf-shaped bract.
Comments & comparison — see notes for genus.
Avistrobus
PINOPSIDA/INCERTAE
d^TRELITZIA 15 (2003)
INCERTAE/PINOPSIDA
pi. 32
Avistrobus foliosus
Birds River
(Bir 1 1 1 Sph 2spp)
all PRE/F/15591b
Holotype
132
c/tRELITZIA 15 (2003)
PINOPSIDA S.V.Meyen 1984
INCERTAE SEDIS order
INCERTAE SEDIS family
HelvetianthllS J.M.And. & H.M.And., gen. nov.
Type species
Helvetianthus tintinnabulum J.M.And. & H.M.And., sp. nov.
Generic diagnosis
A putative pinopsid male cone with simple microsporophylls compris-
ing single, sessile, spherical 4-lobed microsporangiate cupules.
Generic characters
Strobilus: simple, compact, spicate, small (20 X 9 mm); axis relatively
stout (1 mm diam.), free end ca 3 mm, gently curved; microsporo-
phylls spirally attached, sessile, 4 or 5 units per gyre.
Microsporophyli. spherical cupulate heads (ca 3 mm diam.), characteristi-
cally dehiscing along 4 clear suture lines into 4 lobes; leathery; surface
ornamentation clear, isodiametric cells aligned and curving.
Microsporangium: no separate microsporangia observed.
Pollen: disaccate grains, cf Alisporites.
Etymology
Helvetianthus — after Little Switzerland, the type locality.
Global range: 1 sp., Gondwana, Tr. (CRN).
First & last: the single Molteno species described here.
Gondwana Triassic occurrence
SAf— Karoo Basin, 1 TC (6 indivs).
Molteno occurrence
Frequency (F): 1 TC (of 100 sampled in the Molteno).
Diversity (D): 1 species.
Abundance (A): 6 individuals total.
Lit 111: 6 indivs in 550 man-hrs cleaving (1 per ca 10 man-days), extremely rare
Affiliated organs
Female cone— unknown.
Foliage — unknown.
Classification & comparison
Suprageneric classification (family & order incertae)
Prior to isolating clusters of in situ pollen grains from one of the H.
tintinnabulum specimens, we had taken the cupulate heads to be megas-
porophylls and had tentatively placed the strobilus in the Ginkgoopsida. As
a compact male cone with spirally arranged microsporophylls, Hel-
vetianthus appears closer to the Pinopsida or Cycadopsida. In the spherical,
outwardly dehiscing nature of the pollen-bearing heads, however, the genus
is very unlike any other male cone known to belong to either of these class-
es. In that the pollen grains are disaccate, a link with the Pinopsida or
Ginkgoopsida is indicated. Overall, the evidence favours placement in the
Pinopsida, but in a hitherto undefined order and family.
It is interesting to consider that in the angiosperm order Fagales, most
notably in the families Myricaceae, Fagaceae and Casuarinaceae. the male
and female flowers are characteristically distinct and cone-like. The casuar-
inas, in particular, have raised much debate amongst taxonomists as to their
classification. Earlier workers were inclined to the opinion that they were
the most primitive group of angiosperms and probably related to the gym-
nosperms, notably the Pinales and Ephedrales. Later botanists favoured the
view that they were more advanced and that the peculiar features were
extreme specialisations derived in isolation (Hey wood 1993). Recent
cladistic analyses incorporating molecular data (e.g. APG 1998; Nandi et
al. 1998; Savolainen et al. 2000; Soltis et al. 2000) that have revealed con-
siderable changes to our understanding of angiosperm phylogeny, support
the latter view.
H. tintinnabulum
Reconstructions
The four R2-3 reconstructions cover the best of the six specimens of
the reference palaeodeme. They are given the intermediate grade between
R2 and R3 in that the four-lobed spherical heads are rather substantially
reconstructed based on the best-preserved microsporangia in the set of
available individuals.
The composite reconstruction (tf. 1 above) reflects the sum of features
as seen in tfs 2-5 above. These allow a fairly confident reconstruction of
the complete Helvetianthus cone.
Molteno occurrence (elaborated)
For an analysis of the Lit 111 TC. yielding the only known Helveti-
anthus specimens, see And. & And. (in prep.), a sequel to the present work.
Habit & habitat
Like the two previously described ovulate genera, Gypsistrobus and
Avistrobus, which are unique to Aas 41 1 and Bir 111 respectively, Helveti-
anthus is unique to Lit 111. And like those two genera, its great rarity sug-
gests that it is an allochthonous (possibly parautochthonous, considering the
delicate nature of the strobilus) element in the deposit, formed in an
abandoned marginal channel of a braided river with raised banks supporting
dense Dicroidiiim-donmvMcd riverine forest. In view of the nature of the
well-preserved cuticle without stomata it is proposed that Helvetianthus was
borne on an undershrub or herb of the Dicroidium forest.
Evidence for affiliation of organs
Of the seven genera of female fruit and 1 8 of foliage identified at Lit
111, none are evident candidates for linkage with Helvetianthus. While
several of the foliage genera from the TC remain unpreoccupied, there is
no persuasive evidence, from cuticle (And. & And. 1989) or otherwise, for
affiliation. Jeanjacquesia and Moltenia, putative cycadalean fronds (see
Tabs 6a, 13), cannot be excluded from possibility, nor can Saportaea, re-
presented in the Molteno by only one large leaf from Lit 111. A further
possibility is Paraginkgo antarctica, which is placed here in a new genus
as the cuticle is unique and quite unlike that of other Molteno Ginkgoites
species. Of the seven ovulate genera from Lit 111, two remain without pro-
posed male counterparts: Dordrechtites and Fraxinopsis (Tab. 6b). They
offer no clues for pairing with Helvetianthus.
Helvetianthus
PINOPSIDA/INCERTAE
TRELITZIA 15 (2003)
133
Helvetianthus tintinnabulam j.M.And. & h m And., sp nov.
Holotype
Specimen'. PRE/F/22186 a,b; pis 33(1, 2, 6, 7), 34(2).
Assemblage (TC): Lit 111 Dic/Hei; Little Switzerland.
Preservation-, fairly complete strobilus (apical portion missing), part and
counterpart, longitudinal outer view with several cupulate heads; com-
pression in thinly laminated, carbonaceous (good cuticle) dark grey
shale with moderate cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 6 individuals; 2 with counterparts, all are well preserved com-
pressions with cuticle; together they allow the full cone to be recon-
structed.
Sister palaeodemes— nil.
Specific diagnosis— as for genus.
Specific characters— as for genus.
Etymology
tintinnabulum (Lat.) — a bell, with reference to the strobilus appearing like
a string of bells.
Classification & comparison: see notes for genus.
Intactness of cones
All six available cones are fully intact, with the fragile-appearing
cupulate heads preserved essentially in place. Four of these are virtually
complete, while the remaining two are missing the apical part of the cone.
Cuticles (& in situ pollen)
Potential sample: Lit 111,6 indivs.
Macerated (this work): Lit 111, cupulate heads from 5 indivs.
Preservation grade: Grade 5 (excellent), all features clear, large pieces.
Diagnostic characters: cells isodiametric, pentagonal-hexagonal, walls
straight to gently curved; papillae single, bold, central; stomata absent;
other features absent.
In situ pollen: disaccate grains, nonstriate; occur in aggregated masses;
similar to pollen isolated from Pteruchus by Townrow (1962) and dis-
persed grains from Lit 111 identified as Alisporites in And. & And.
(1983, pi. 30, figs 14—16).
Comment: the cuticular structure is visible on the surface of the micro-
sporangia, even at low magnification (pi. 34, figs 1-6).
Significance:
Classification— The cuticle, in its thickness and cellular features, is
most reminiscent of the ginkgoopsid genera (e.g. Dicroidium) which would
suggest placing Helvetianthus in this class. The disaccate grains would tend
to support this. In view of its (remotely) cone-like appearance, however, we
have placed it in the pinopsids. Helvetianthus remains a phylogenetic enigma.
Affiliations— We have not proposed any likely affiliates for Helveti-
anthus. If the genus should prove to be ginkgoopsid. then it might conceiv-
ably affiliate with Paraginkgo (found most commonly at Lit 111) which
remains without male or female counterparts. But the cuticle of these two
genera does not corroborate this.
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
o
CO
<
INCERTAE/PINOPSIDA
Helvetianthus
d/TRELITZIA 15 (2003)
Holoytpe
Little Switzerland
(Lit 1 1 1 Dic/Hei)
PRE/F/22186a
Helvetianthus tintinnabulum
pi. 33
PINOPSIDA/INCERTAE
135
^VtRELITZIA 15 (2003)
Little Switzerland
INCERTAE/PINOPSIDA
pi. 34
Helvetianthus tintinnabulum
136
TRELITZIA 15 (2003)
CYCADOPSIDA P.D.W.Bamard & A. G. Long 1975
CYC AD ALES Engl. 1892
INCERTAE SEDIS family
Androstrobus Schimp. 1870
Type species:
Androstrobus zamioides Saporta, in Schimper ( 1870). Etrochey, France;
Bathonian, Jurassic.
Generic concept— see Saporta, in Schimper (1872, p. 199)
A cycadopsid male cone of small size, cylindrical to oval shape, and
with microsporophylls bearing sessile microsporangia on abaxial surface.
Generic characters (based on the two Molteno species)
Strobilus : simple compact cone, relatively small (17-35 mm long), oval to
narrowly oblong; axis stout, strongly flexed proximally, free end to 17
mm; microsporophylls helically attached, in 4 to 13 gyres of 8-10
units.
Microsporophyll : scale variously shield-shaped, apex rounded to mucro-
nate. margins entire, face concentrically ornamented.
Microsporangium : unknown.
Etymology
Androstrobus— Andros (Gr.), male; strobilos (Gr.), cone.
Global range: numerous spp., Pangaea, U.Tr.-L.K.
Gondwana Triassic occurrence
SAf— Karoo Basin, 2 TCs (2 indivs).
Molteno occurrence
Frequency (F): 2 TCs (of 100 sampled in the Molteno).
Diversity (D): 2 species.
Abundance (A): 2 individuals total, very rare.
Kra 311 Die odo: 1 indiv. in 13 man-hrs cleaving (1 per 1 man-day) very rare
Pen 321 Dic/Ris: 1 ” ” 35 ” ” (1 ” 3 ” ) ”
The great scarcity of these cones follows the same pattern witnessed
for the voltzialean male cones.
Affiliated organs (Molteno)
Female cone : unknown.
Foliage: Pseudoctenis — Grade 2 (Kin. reinf. ).
Were the only information available that from the Molteno, it is doubt-
ful that the affiliation between Androstrobus and Pseudoctenis would be
suggested. Pseudoctenis (Tab. 6a), by far the most frequent and abundant
of the four cycad foliage genera recognised in the Molteno, occurs in 21
TCs. In seven of these TCs it is a common (>1%) to co-dominant (>20%)
element of the assemblage, yet in none does Androstrobus appear— an
anomaly (assuming the affiliation is correct) ascribable to some undefined
combination of seasonality and taphonomy (see also Tab. 45, p. 183).
Classification & comparison
Suprageneric classification (Incertae sedis/Cycadales)
Harris (1964) entertained little doubt that his Androstrobus specimens
from the Yorkshire Jurassic represent cycads. He recognised three distinct
species of these cones and affiliated them with the frond genera Nilssonia,
Pseudoctenis and Ctenis, respectively, with varying degrees of confidence.
Nor did Harris doubt that certain other species of Androstrobus from the
Upper Triassic to Cretaceous of Europe and Russia were true cycad male
cones. Hill ( 1990). through his work on the ultrastructure of Androstrobus
balmei , verified the link to cycads.
Neither of the Molteno specimens is sufficiently well preserved to
allow certain comparison with the Yorkshire or other Androstrobus mater-
ial. Nor do either yield cuticle or in situ pollen, yet their general appearance
is very like the Laurasian genus.
It is quite possible, though, that one or other of the Molteno specimens
could be included in Switzianthus (Ginkgoopsida), or even amongst the
pinopsids. The latter option seems less likely in view of the fact that we
already recognise at least five genera of male conifer cones in the Molteno,
but only four foliage genera.
Intactness of cones
As noted above, the two Molteno species, each based on a single cone,
are known only in their intact state (tfs 1-6 opposite). The microsporangia
remain unknown.
Reconstructions (Molteno species)
In the reconstructions of the two Androstrobus species from the
Molteno, little is left to interpretation— at least as far as the general outer
form of the cone is concerned. The holotype (only specimen) for each
species is more or less complete— apart from the pedicel of A. kraaiovalis ,
which is conjecture.
Aside from their distal laminae, on the other hand, the morphology of
the microsporophylls cannot be reconstructed and the microsporangia
remain unknown.
Adaptive radiation (Molteno diversity)
The two Molteno species identified here are markedly different and
distinguished according to overall shape and size of strobilus along with
scale morphology. They are based on the following TCs/reference
palaeodemes, which derive from the same open-woodland habitat, but from
different levels within the stratigraphic sequence.
A. peninsiformis— Pen 321 Dic/Ris (Peninsula), 1 indiv.
Dicroidium open woodland; Cycle 2f (Indvve Member)
A. kraaiovalis— Kra 311 Die odo (Kraai River), 1 indiv.
Dicroidium open woodland; Cycle 3 (Mayaputi Member)
Androstrobus
CYCADALES
TRELITZI A 15 (2003)
137
Aiidrostrobus peninsiformis J.M.And. & H.M.And.. sp. nov.
Holotype
Specimen: PRE/F/13687a,b; pi. 35(1-5).
Assemblage (TC): Pen 321 Dic/Ris; Peninsula.
Preservation: virtually complete cone (apex uncertain), part and counter-
part, longitudinal outer view with overlapping scales intact; 3D mould
and cast in thickly laminated, medium light grey shale with poor cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimen: 1 complete indiv.
Sister palaeodemes — nil.
Specific diagnosis
An Androstrobus species with long cylindrical strobilus and acute-
tipped, weakly ornamented microsporophylls.
Specific characters
Strobilus: narrowly oblong (38 X 8 mm).
Microsporophyll: distal lamina with mucronate apex and relatively ill-
defined concentric pattern.
Etymology
peninsiformis— after the type locality. Peninsula, and the shape of the cone.
Comment & comparison— see notes for genus.
Androstrobus kraaiovalis J.M.And. & H.M.And., sp. nov.
Holotype
Specimen PRE/F/8231a,b; pi. 36(1-6).
Assemblage (TC): Kra 311 Die odo; Kraai River.
Preservation: virtually complete cone (free axis missing), part and counter-
part. longitudinal outer view with overlapping scales intact; impres-
sion, clearly preserved; in grey, irregularly bedded shale.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimen: 1 intact indiv.
Sister palaeodemes— nil.
Specific diagnosis
An Androstrobus species with short oval strobilus and obtuse-tipped,
strongly ornamented microsporophylls.
Specific characters
Strobilus: oval (20 X 11 mm).
Microsporophyll: distal lamina with rounded apex and sharply defined
concentric pattern.
Etymology
kraaiovalis— after the type locality Kraai River, and the oval shape of the
cone.
Comment & comparison — see notes for genus.
CYCADALES
Androstrobus
138
^/tRELITZIA 15 (2003)
Peninsula
(Pen 321 Dic/Ris)
all PRE/F/13687a
Holotype
CYCADALES
Androstrobus peninsiformis
CYCADALES
pi. 36
Androstrobus kraaiovalis
140
d^TRELITZIA 15 (2003)
CYCADOPSIDA P.D. W. Barnard & A. G. Long 1975
CYC AD ALES Engl. 1892
INCERTAE SEDIS family
Pseudoctenis Seward 1911
Type species
Pseudoctenis eathiensis (Richards) Seward 1911.
Upper Oolite, Scotland; Jurassic.
Generic concept (for Gondwana Triassic only)
A cycadopsid frond bearing nondehiscing pinnae with entire margins
and parallel veins that fork once or twice at or near the base.
Generic characters (after Harris 1964)
Leaf. 'Leaf large, elongated, simply pinnate. Pinnae broad or narrow and
elongated, lanceolate or parallel-sided, arising laterally on the rachis.
Pinna margins entire, apex truncate or contracted, base expanded or
contracted; veins numerous, parallel, simple or forked, not anastomos-
ing.’
Cuticle : see And. & And. (1989. p. 281); this vol., tfs 1,2 below.
Etymology
Pseudoctenis— pseudes (Gr. ), false; denis, a fossil cycad genus.
Global range: numerous spp., Pangaea, U. Tr.-U. K.
Gondwana Triassic occurrence
Diversity (D): 11 foliage species.
Ubiquity (U): 3 continents (of 5 comprising Gondwana).
Frequency (F); 16 degree squares (of the 84 across Gondwana).
Abundance (A): 3% (the norm in Molteno TCs).
Longevity (L): 14 myrs (Lower Anisian to Lower Camian).
Colonisation success: FUDAL rating 16/3/11/3/14 = 47.
Intermediate success (Grade 3); Pseudoctenis was the ninth most
prominent genus in the Gondwana Triassic; it was diverse, but of only
moderate ubiquity, frequency, abundance and longevity.
Endemism: of the 11 Gondwana Triassic species, six occur more or less
widely through the realm, 1 is a basin endemic, and four are single-
assemblage endemics.
Molteno occurrence
Frequency (F): 21 TCs (of 100 sampled in the Molteno).
Diversity (D): 9 species.
Abundance (A): co-dominant (21-25%) in 2 TCs; occasional to common
(1-3%) in 2 TCs; and <1% in the other 17 TCs.
Habit: probably a cycad-like plant.
Preferred habitat: The genus occurs mostly in high diversity assemblages
that are generally Dicroidium-dom'mated and probably represent forest
and woodland associations of the riverbank (levee) and other relative-
ly elevated ground.
Affiliated organs
Female strobilus: unknown
Male strobilus: Androstrobus — Grade 2 (Kin. reinf.); see note under Andro-
strobus (p. 136).
Classification & comparison (adapted from And. & And. 1989, p. 280)
Intergeneric comparisons
Gondwana Triassic cycadalean genera— The four genera ( Pseudoc-
tenis, Jeanjacquesia, Ctenis and Moltenia) of Molteno cycads are clearly
distinguished both on the basis of mega- and micromorphological charac-
ters. The cuticle, in particular, provides an indication of the taxonomic dis-
tance between the genera. They are considered here as probably represent-
ing four distinct families (p. 54). The reproductive organs remain unknown,
thus shedding no light on the matter.
Other cycadalean genera— We have elected not to employ the genus
Nilssonia which has been shown by Meyen (1984) to be a form-genus af-
filiating with distinctly different types of female reproductive structures. In
our Gondwana Triassic hypodigms (And. & And. 1989) entire fronds that
might have been included as Nilssonia are listed under Taeniopteris and
segmented fronds under Pseudoctenis.
Bennettitalean (cycadeoid) genera— Pseudoctenis is distinguished
from the bennettitalean genus Pterophyllum solely on the basis of its cuticle.
It must be noted, though, that for the great majority of the Molteno and
Gondwana Triassic fronds identified as Pseudoctenis there exists no pre-
served cuticle.
Interspecific comparisons
We assumed in And. & And. ( 1989) that the greater proportion of the
1 1 Gondwana Triassic Pseudoctenis species recognised were closely related
within the scope of a natural genus. In view, however, of the paucity of
cuticle (poor material from only two of the nine Molteno species has been
obtained) and lack of affiliated fruit, there remains no way of confirming
this view.
The nine Pseudoctenis species from the Molteno are quite distinctive
when only their reference palaeodemes are considered, but the position
becomes far less clear when all 25 palaeodemes from 1 5 assemblages are
taken into account. The morphological ranges seen in the different
palaeodemes overlap in a complex fashion: rarely do those of any two
palaeodemes approximately coincide.
P. harringtoniana
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
Pseudoctenis
CYCADALES
d^TRELITZIA 15 (2003)
141
CYCADALES
Pseudoctenis
142
d^TRELITZIA 15 (2003)
CYCADOPSIDA P.D.W.Bamard & A. G. Long 1975
CYCADALES Engl. 1892
INCERTAE SEDIS family
Jeanjacquesia J.M.And. & H.M.And. 1989
Type species
Jeanjacquesia switzipinnata J.M.And. & H.M.And. 1989.
Little Switzerland, Karoo Basin, S. Africa; Camian, Triassic.
Generic diagnosis
A cycadopsid frond bearing readily dehiscing pinnae with entire margins
and parallel veins that fork only once at or near the base.
Generic characters [based on J. switzipinnata & J. spA (Bir 111)].
Leaf, relatively large, shape unknown, simply pinnate; pinnae readily de-
tach (attachment unknown), margins entire, linear oblong, apex
unknown, base contracted, asymmetrical; veins moderately to relative-
ly well spaced, parallel and not meeting lamina margin, simple or
forked near base, not anastomosing.
Cuticle: see And. & And. (1989, p. 337); this vol., tfs. 3-6 opposite.
Eponymy
Jeanjacquesia— after Jean-Jacques Rousseau, French philosopher.
Global range: 3 spp., Gondwana, Tr. (ANI-NOR).
First & Last : the three Molteno species.
Gondwana Triassic occurrence
Frequency (F): 3 degree squares (of the 84 across Gondwana).
Ubiquity (U): 1 continent (of 5 comprising Gondwana).
Diversity (D): 3 foliage species.
Abundance (A): <1% (the norm in Molteno TCs).
Longevity (L): 2 myrs (Camian).
Colonisation success: FUDAL rating 3/1/3/-/2 = 9.
Minimum success (Grade 1 ): Jeanjacquesia was the 23rd most promi-
nent genus in the Gondwana Triassic; it was of very low frequency,
ubiquity, diversity, abundance and longevity.
Endemism: all three species are single-assemblage endemics.
Molteno occurrence
Frequency (F): 3 TCs (of 100 sampled in the Molteno).
Diversity (D): 3 species.
Abundance (A): extremely to vanishingly rare (<1%) in the 3 TCs.
Habit: probably a cycad-like plant.
Preferred habitat: the type species, J. switzipinnata , is a very rare element
(six individuals collected) in the Lit 111 Dic/Hei assemblage, which is
dominated in particular by Dicroidium odontopteroides. It most likely
occupied the understorey of the riverine forest.
Affiliated organs — unknown.
Classification & comparison (adapted from And. & And. 1989, p. 336)
Intergeneric comparisons
Gondwana Triassic cycadalean genera— Jeanjacquesia, on the basis
of both general morphology and cuticle, is quite distinct from the other four
Gondwana Triassic cycad genera. It is most like Pseudoctenis, but the lat-
ter differs in having pinnae with symmetrical bases that do not dehisce.
Other cycadalean genera— Jeanjacquesia does not exhibit the heavily
developed pinna base usually seen in the extant cycad genera (e.g.
Encephalartos, Zarnia) that occasionally drop their pinnae. J. switzipinnata
does have typically cycadaceous cuticle, which, in its perfectly hypostom-
atic nature, its well developed guard cells, cutinised subsidiary cells and
strongly curved cell walls, particularly resembles Zamia (Greguss 1968).
Its cuticle shows less similarity to any of the Yorkshire Jurassic cycad gen-
era.
Bennettitalean (cycadeoid) genera— Zamites and Otozamites are the
cycadeoid genera most similar to Jeanjacquesia, but Zamites has an essen-
tially symmetrical base, while that of Otozamites is strongly asymmetrical,
with the upper angle developed into an auricle. Both have somewhat
spreading venation and both have typically cycadeoid cuticle.
Interspecific comparisons
The two species, J. switzipinnata and J. spA, are sufficiently similar to
appear to belong to a single natural genus, but since cuticle is known only
for the former and fruit for neither, this cannot be confirmed.
J. spB, based on a single Molteno (Maclear) specimen in Du Toit’s col-
lections (Du Toit 1927), was placed in Jeanjacquesia for convenience in
And. & And. (1989), but might equally tentatively have been included in
Zamites (cycadeoid), Pseudoctenis (cycad), or some unnamed genus. It is
not illustrated in the present volume.
Each species is represented solely by the reference palaeodeme.
GONDWANA TRIASSIC, ‘GEOSTRAT DISTRIBUTION
Jeanjacquesia
CYCADALES
d^TRELITZIA 15 (2003)
143
The Cycadales at family level
The extant cycads are included in three families (Stevenson 1990,
1992):
Cycadaceae— 1 genus ( Cycas ), 14 species.
Stangeriaceae— 1 genus ( Stangeria ), 2 species.
Zamiaceae — 8 genera ( Encephalartos , Zamia etc.), ca 110 species.
In an attempt to resolve, however imperfectly, the possible family-
level taxonomy of the four Molteno cycad foliage genera, we are limited
to a consideration of their general morphology and cuticular features.
Female cones from the formation (and, indeed, the entire Gondwana
Triassic) remain unknown, while supposed male cones are represented by
only two individuals that are very uncertainly affiliated with the foliage
genus Pseudoctenis.
Though exceptions must be acknowledged, and though diagnosed on
the basis of their female cones, the three extant families can be separated
reasonably successfully on consideration of their foliage morphology and
cuticles alone (Greguss 1968). The Cycadaceae (Cycas) species all have
linear leaves with a single distinct vein or midrib and cuticles showing
large pits along the cell walls; the Stangeriaceae ( Stangeria ) species have
taeniopteroid leaves and cuticles with meandering cell walls; while the
Zamiaceae sport parallel-veined leaves and cuticles with gently curved to
sinuous cell walls.
In the light of the above, it seems that the most likely option is to con-
sider each of the four Molteno genera to represent a different cycad fam-
ily within a vigorously sprouting cycadopsid phylogenetic tree following
the end-Permian extinction. A comparative study of their foliage mor-
phology and cuticles, outlined below, suggests that they are as distinct
from one another as are the core genera characterising the extant families:
Pseudoctenis (9 Molteno species)
Leaves— pinnae not dehiscing, entire, with parallel venation.
Cuticle— cell walls relatively straight, subsidiary cells apparently undif-
ferentiated.
Moltenia (4 Molteno species)
Leaves— pinnae not dehiscing, toothed, with parallel venation.
Cuticle— cell walls, strongly undulating, subsidiary cells apparently
undifferentiated.
Jeanjacquesia (3 Molteno species)
Leaves— pinnae dehiscing, entire, with parallel venation.
Cuticle— cell walls distinctly sinuous, subsidiary cells highly developed
and well-cutinised.
upper lower BP/2/1809
1
1
'll ll 1
\
x ' , '
' ! 1
' - '
1
1 II
j
ill"
' '«
Ctenis (2 Molteno species)
Leaves— pinnae dehiscing or not dehiscing, margins lobed or irregular,
with clearly anastomosing venation.
Cuticle— cell walls strongly sinuous to meandering (stomata unknown).
Alternative suprageneric taxonomic options quite obviously exist, but
short of well-affiliated fruiting material no certain conclusions are possible.
HMA, for instance, feels that the available evidence is insufficient to dif-
ferentiate families.
tfs 1-6 from And. & And., 1989
CYCADALES
Jeanjacquesia
144
Cff TRELITZIA 15 (2003)
CYCADOPSIDA P.D.W. Barnard & A. G. Long 1975
CYCADALES Engl. 1892
INCERTAE SEDIS family
Cteilis Lindl. & Hutton 1834
Type species
Ctenis falcata Lindl. & Hutton 1834.
Yorkshire, England; Jurassic.
Generic concept (for Gondwana Triassic only)
A cycadopsid frond bearing essentially nondehiscing pinnae with entire
to irregular margins and parallel veins that fork and clearly anastomose
throughout the lamina length.
Generic characters (adapted from Harris 1964)
Leaf, shape and size not given, once pinnate; pinnae usually with entire
margins, inserted laterally on the rachis, shape (including apex and
base) and base not given; veins several, more or less parallel, anasto-
mosing, meeting lamina margin.
Cuticle', see And. & And. (1989, p. 342); this vol., tfs 3, 4 opposite.
Etymology
Ctenis— Ktenos (Gr.), comb.
Global range: numerous spp., Pangaea, Tr.-K.
Gondwana Triassic occurrence
Frequency (F): 4 degree squares (of the 84 across Gondwana).
Ubiquity (U): 3 continents (of 5 comprising Gondwana).
Diversity (D): 2 foliage species.
Abundance (A): <1% (the norm in Molteno TCs).
Longevity (L): 2 myrs (Lower Camian).
Colonisation success : FUDAL rating 4/3/2/— /2 =11.
Minimum success (Grade 1): Ctenis was the 22nd most prominent
genus in the Gondwana Triassic; it was fairly ubiquitous and of very
low frequency, diversity, abundance and longevity.
Endemism', the two Molteno species are single-assemblage endemics (the
non-Molteno specimens are not identifiable to species).
Molteno occurrence
Frequency (F): 2 TCs (of 100 sampled in the Molteno).
Diversity (D): 2 species.
Abundance (A): very rare to vanishingly rare (<1%) in the 2 TCs.
Habit', probably a cycad-like plant.
Preferred habitat : The two Ctenis species from the Molteno Fm. occur in
high diversity assemblages: the one being very rare from Kon 211/221
dominated by Asterotheca and Pseudoctenis , the other vanishingly rare
from Umk 1 1 1 dominated by Dicroidium odontopteroides.
Affiliated organs— unknown.
Classification & comparison (adapted from And. & And. 1989, p. 342)
Intergeneric comparisons
Gondwana Triassic cycadalean genera— Ctenis, on frond morphology,
is readily distinguished from the other four Gondwana Triassic cycad gen-
era. There is no particular genus that it resembles most closely.
Other cycadalean genera— We are unaware of any other cycad genera,
extinct or extant, in which anastomosing venation is a diagnostic charac-
teristic. It should be noted, though, that the extant genus Stangeria (with a
single species) and at least two of the 30 species of the extant
Encephalartos show venation with occasional anastomoses.
Bennettitalean ( cycadeoid ) genera— Dictyozamites is the only
cycadeoid genus with anastomosing venation, but it is otherwise quite
unlike Ctenis in that the pinnae are strongly contracted at the base (which
is asymmetrical) leaving only a narrow section attaching to the rhachis.
Interspecific comparisons
The two species from the Molteno identified as belonging to the pan-
demic genus Ctenis may be quite unrelated. C. biloba has produced tiny
fragments of cuticle which do not exclude identity with Ctenis. Neither
species has affiliated fruit.
The reference palaeodemes of the two species, consisting of only one
and two individuals respectively, are distinctly dissimilar. No additional
palaeodemes occur.
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
Ctenis
CYCADALES
d^TRELITZIA 15 (2003)
145
CYCADALES
Ctenis
146
d/TRELITZIA 15 (2003)
CYCADOPSIDA P.D.W.Bamard & A. G. Long 1975
CYC AD ALES Engl. 1892
INCERTAE SEDIS family
Moltenia A.L.duToit
Type species
Moltenia dentata A.L.du Toit 1927.
Umkomaas Valley, Karoo Basin, S. Africa; Camian, Triassic.
Generic diagnosis
Acycadopsid frond bearing nondehiscing pinnae with toothed margins
and parallel veins that fork in the proximal half to two-thirds of the lamina.
Generic characters (based on the four Gondw. Trias, species)
Leaf, small to large, shape uncertain (apparently narrowly ovate in one spe-
cies), once pinnate; pinnae laterally attached, margins variously
toothed, narrowly elliptic, apex usually irregularly toothed, base
markedly contracted; veins closely (1 spp.) to well spaced (3 spp.),
spreading to meet lamina margin at each tooth, repeatedly forking but
not anastomosing.
Cuticle', see And. & And. (1989, p.348); this vol., tf. 2 opposite.
Etymology
Moltenia— named by Du Toit in reference to the Molteno Fm.
Global range: 4 spp., Gondwana, M.-U.Tr. (LAD-CRN).
First. Moltenia ( Pseudoctenis wardii) (Artabe 1985); Los Menucos Fm.,
southern Argentina, South America.
Last : the three Molteno species (And. & And. 1989).
Gondwana Triassic occurrence
Frequency (F): 4 degree squares (of the 84 across Gondwana).
Ubiquity (U): 3 continents (of 5 comprising Gondwana).
Diversity (D): 4 foliage species.
Abundance (A): <1% (the norm in Molteno TCs).
Longevity (L): 4 myrs (Upper Ladinian to Lower Camian).
Colonisation success : FUDAL rating 4/3/4/— /4 = 15.
Limited success (Grade 2): Moltenia was the 20th most prominent
genus in the Gondwana Triassic; it was fairly ubiquitous but of very
low frequency, diversity, abundance and longevity.
Endemism: of the four Gondwana Triassic species, one is known from two
continents, one is a single-basin endemic, and two are single-assem-
blage endemics.
Molteno occurrence
Frequency (F): 5 TCs (of 100 sampled in the Molteno).
Diversity (D): 4 species.
Abundance (A): very rare to vanishingly rare (<1%) in all 5 TCs.
Habit : probably a cycad-like plant.
Preferred habitat : The genus occurs as a particularly rare component in
high to very high diversity assemblages. Those assemblages from
Hlatimbe, Umkomaas and Little Switzerland, in which it occurs, are all
dominated by Dicroidium and in three of five cases by D. odon-
topteroides. All four species of Moltenia most likely occupied the
understorey of riverine forest.
Classification & comparison (adapted from And. & And. 1989, p. 346)
Intergeneric comparisons
Gondwana Triassic cycadalean genera— Moltenia, on both general
frond morphology and cuticle, is readily distinguished from the other four
Gondwana Triassic cycad genera. It is most like Pseudoctenis , but the latter
differs in the pinnae having entire margins and parallel venation, and in the
epidermal cell walls being straight rather than strongly sinuous.
Other cycadalean genera— In the toothed nature of the pinna margin,
Moltenia is very like the extant South African cycad genus Encephalartos,
but in the attachment of the pinnae it is markedly different. The epidermal
cell walls of Encephalartos are gently curved, not strongly sinuous as in
Moltenia.
Bennettitalean (cycadeoid) genera— Moltenia, in general frond mor-
phology, does not resemble any of the cycadeoids. While it does display
sinuous cell outlines, these do not approach the typical close meandering as
witnessed in the cycadeoids; nor do any other typical cuticular features of
the latter order appear.
Interspecific comparisons
The four species comprising Moltenia give the clear impression of repre-
senting a single natural genus, but this cannot be confirmed either on the
basis of cuticle (known for only one species) or affiliated cones (unknown).
The reference palaeodemes (RPs) of the four species are markedly dis-
tinct. The three additional palaeodemes (sister palaeodemes, SPs), repre-
senting two of the species, are readily compared to the relevant RPs, hence
supporting rather than confusing the recognition of species.
Species
Ref. Pal.
indivs
(in RPs)
indivs
(in SPs)
M. dentata
Umk 111
18
1 (Lit 111)
M. paucidentata
”
1
-
M. feistmantelii
Hla 213
5
-
M. gracilidentata
1
6 (Hla 211, Hla 212)
Moltenia species & abundance, Molteno Fm.
Affiliated organs — unknown.
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
Moltenia
CYCADALES
d^TRELITZIA 15 (2003)
147
CYCADALES
Moltenia
148
d^TRELITZIA 15 (2003)
GINKGOOPSIDA S.V.Meyen 1987
PELTASPERMALES F.Nemejc 1968
PELTASPERMACEAE H.H.Thomas ex T.M. Harris 1937
Peltaspermum T.M.Harris 1937
Synonym: Meyenopteris R.J.Poort & J.H.F.Kerp 1990.
Type species
Peltaspermum rotula T.M. Harris 1937.
Tancredia River ( Lepidopteris Bed), Scoresby Sound, E. Greenland;
Rhaetic, U. Triassic.
Generic concept
A ginkgoopsid strobilus of linear-cylindrical shape with megasporo-
phylls of one to five peltate multilobed discs bearing abaxial ovules.
Generic characters (Molteno Fm.)
Strobilus: simple, racemose, compact, radially symmetrical, of medium size
( ca 100-175 mm long); axis relatively gracile, generally slightly
curved; megasporophylls numerous, helically arranged.
Megasporophyll: simple to compound, pedunculate; ovuliferous discs
peltate (4-8 mm diam.), radially symmetrical, with or without distinct
apical cap. multilobed (6-12 lobes); ovules/seeds abaxial, naked, pen-
dent, 1 per lobe.
Ovule/seed: tetrahedral (2 X 1 mm), unwinged, with bifid micropyle.
Etymology
Peltaspermum— pelt a (Lat.), small shield; sperma (Gr.), seed, with refer-
ence to the peltate ovuliferous discs.
Global range: numerous spp., Pangaea, U.P.-L.K.
Gondwana Triassic occurrence
SAm— N. Argentina, 2 TCs (2 indivs).
SAf— Karoo Basin, 17 TCs (>200 indivs).
Aus— New South Wales & Victoria, 3 TCs (3 indivs).
Molteno occurrence
Frequency (F): 17 TCs (of 100 sampled in the Molteno).
Diversity (D): 4 species.
Abundance (A): 257 individuals total, rare to extremely rare in top 8 TCs.
Kon 222 Die odo: 24 indivs in 40 man-hrs (6 per 1 man-day) rare
Aas 411 Dic/Sph:
88
” ”512 man-hrs
(1-2 per 1
man-day) very rare
Bir 111 Sph 2spp:
67
” ” 550
(1-2 ” 1
” ) ”
Tel 111 Heielo:
14
” ” 90
(1-2 ” 1
., ) .,
Maz 211 Hei/Dic:
9
” ” 85
(1 ” 1
” ) ”
Mat 111 Die dub:
4
” ” 65
(1 ” 1
” ) ”
Umk 111 Die 2spp:
17
” ” 400
(1 ” 3
” ) ”
Lit 111 Dic/Hei:
11
” ” 550
(1 ” 5
” ) extr. rare
Peltaspermum is very much more common in the two lake deposits
(Aas 41 1 and Bir 111) than in the Dicroidium riparian forest habitats (Umk
111 and Lit 111).
Affiliated organs
Male strobilus: Antevsia— Grade 3 (Kin. reinf., Mut. occ.).
Foliage: Lepidopteris— Grade 4 (Mor. corr.. Cut. corn).
Classification & comparison
Suprageneric classification (Peltaspermaceae/Peltaspermales)
The prominent early Mesozoic pteridosperms, including Peltasper-
mum, Umkomasia (corystosperm) and Caytonia, have been very variously
classified. Crane (1985, 1988), on the basis of cladistic analysis, found the
corystosperms more closely related to the glossopterids and Caytonia than
to the peltasperms. Meyen (1987) included the families Umkomasiaceae
(= Corystospermaceae) and Peltaspermaceae within the order Peltasper-
males, and identified the Caytoniales as a separate order. Stewart & Rothwell
(1993) included the families Corystospermaceae and Peltaspermaceae in the
order Caytoniales. Taylor & Taylor (1993) treated the Corystospermales,
Peltaspermales and Caytoniales as separate orders. In And. & And. (1989,
pp. 64—67), we followed Meyen’s classification, finding the cuticle of the
corystosperm and peltasperm foliage compellingly similar.
In the light of our present study, we consider the Umkomasia and
Peltaspermum strobili clearly distinct at order level — in line with Taylor &
Taylor (1993)— and we place the two orders in the class Ginkgoopsida.
Intergeneric comparison (Gondwana Triassic)
For discussion, see under Matatiella (p. 172).
Peltaspermum
PELTASPERMALES
TRELITZIA 15 (2003)
149
Reconstructions
The reconstructions of the four of the five Molteno Peltaspermum
species that occur in our collection are based largely on their respective
holotypes. In contrast to many other ovuliferous genera such as the ben-
nettitopsids Fredlindia and Lindtheca, relatively little interpretation has
been necessary. Peltaspermum is already well known through studies by
Harris (1932a) and Townrow (1960) and excellent specimens are at hand.
We have assumed a gently arching, pendulous form of the strobilus for
three of the species and an erect mode for P. tridiscum , with its robust, zig-
zag axis.
Evidence for affiliation of organs
The affiliation between Peltaspermum and Lepidopteris has long been
accepted without question — supported by numerous instances of mutual
occurrence— throughout their range in Gondwana and Laurasia.
The link is confirmed by Molteno data (Tab. 40), although there are
interesting discrepancies in occurrence patterns. All but one of the 17
Peltaspermum- yielding TCs also yield Lepidopteris (known from 30 of the
100 Molteno TCs); yet Peltaspermum occurs in only one of the five
Molteno TCs (Aas 411) where Lepidopteris exceeds 1% of the foliage
assemblage.
Peltaspermales beyond Gondwana Triassic
Laurasian Rhaeto-Liassic — Peltaspermum , with its familiar af-
filiates Antevsia and Lepidopteris, occurs widely (e.g. Greenland,
Germany, Sweden, Russia) in the Rhaeto-Liassic of Laurasia. It was
first recorded by Harris (1932a) and later described in Harris (1937).
Laurasian Permian— Peltaspermum-Uke megasporophylls occur
in the Upper Permian Tatarian- Flora of the Russian Platform, where
they are placed in separate genera such as Peltaspermopsis and
Lopadiangium (Gomankov & Meyen 1986). Some of these are affil-
iated with leaves, e.g. Tatarina, which are very different from Lepi-
dopteris. Peltaspermum megasporophylls from the Zechstein Flora
of Western and Central Europe were revised by Poort & Kerp ( 1990)
and affiliated with Lepidopteris (Callipteris)- like leaves.
Molteno occurrence (elaborated)
The five Molteno Peltaspermum species recognised here are very dis-
tinctive and display an interesting pattern of distribution in the formation.
Only P. monodiscum is widespread, while the remaining taxa appear
restricted to particular habitats. This type of pattern, with one possibly ple-
siomorphic (classical) species of widespread occurrence spawning several
derived, apomorphic (mannerist) species of restricted occurrence, is seen
also in other gymnospermous reproductive genera (e.g. Telemachus,
Dordrechtites, Fraxinopsis) and foliage genera (e.g. Ginkgoites) in the
Molteno.
P. thomasii— occurs only at Umk 111 (riparian forest) where it was collected by
Thomas (1933) and Townrow (1960). We have found no additional material that we
can attribute to this species.
P. monodiscum— occurs in 10 of the 17 TCs yielding Peltaspermum and would appear
to have flourished in or on the periphery of a range of habitats; Dicroidium riparian
forest (mature and immature), Dicroidium open woodland, Sphenobaiera closed
woodland and Heidiphyllum thicket.
P. tridiscum— occurs in three TCs, all representing Dicroidium open woodland and all
in the extensive cherty horizon of the Konings Kroon/Peninsula district.
P. turbanatum—a\so occurs in three TCs (notably Bir 111 and Aas 411). but in this
case all are floodplain lake deposits representing Sphenobaiera closed woodland.
Although the species is known only from these deposits, it clearly extended through-
out the duration of the Molteno with Aas 411 occurring low in the formation (Cycle
1) and Bir 111 in the upper levels (Cycle 5).
P. quindiscum — '\s confined to four TCs representing riparian forests (mature and
immature) all occurring low in the Indwe Member (Cycle 2) of the formation.
Gondwana Triassic occurrence (elaborated)
While Peltaspermum is relatively frequent and common in the
Molteno and the affiliated foliage (Lepidopteris) occurs commonly and
widespread throughout Gondwana, the ovulate organ remains enigmatical-
ly rare outside South Africa. Only three records (very fragmentary materi-
al) from Australia and two from South America are known. This extreme
rarity is difficult to explain through the lack of intensive or extensive sam-
pling alone.
South America
Peltaspermum was first recorded from South America by Baldoni &
De Cabrera (1977), who illustrated a single disc from the Portezuelo Fm.,
and subsequently by Zamuner et al. (1999), who figured a fragmentary
strobilus from the Cortaderita Fm. Neither specimen is sufficiently pre-
served for identification to species.
Australia
The three records of Peltaspermum from Australia are all of single dis-
sociated megasporophyll discs. They are from three well-separated deposi-
tional areas: Bald Hill in Victoria (Douglas 1973); the Lome Basin in New
South Wales (Holmes & Ash 1979); and the Dubbo district of the Great
Artesian Basin also in New South Wales (Holmes 1982). The material is
inadequate for specific identification.
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
PELTASPERMALES
Peltaspermum
150
d/TRELITZIA 15 (2003)
Tab. 39.
PELTASPERMUM HYPODIGM, Gondwana Triassic occurrence
Species
Intact-
ness
Molteno
P. thomasii
P. monodiscum
P. tridiscum
P. turbanatum
P. quindiscum
P. spp. indet
Intact strobili
Fragmentary ”
Isolated megasp.
AUTHOR
SUBREGION
FORMATION
LOCALITY
NAME
Indivs ILLUSTRATION
SOUTH AMERICA
s
1977
Baldoni & De C.
Barreal NA2
25 Portezuelo Fm
Ao. Panul
Antevsia sp.
1
pi 1(1)
-
-j -j - -
1
-N i
1999
Zamuner et at.
23 Cortaderita
Level 2
Peltaspermum sp.
1
pl 4(C,D)
-
-I 'I "I -
1
- i -
SOUTH AFRICA
1933
Thomas
Elliot T Ka9
24 1 Molteno
Konings Kroon
Pteruchus edwardsi
1
tf 42, pl 24(74)
- - - -
1
i -T-
Underberg Ka4
Umkomaas
Lepidopteris natalensis
1
tf 55, pl 24(78)
1
-MH -
-
ihdr
1955
” (specimen repeated from Thomas 1933)
*
tf 1 A
*
-i d .
-
*1-1-
1960
Townrow
Underberg: Ka4
24 Molteno
Umkomaas
Peltaspermum thomasi
3
tflO(B-F), 11(A,B), pl 58(2,8)
3
- - - -
-
3; 1 -
1978-1999: And. & And. Molteno literature not included in this table
1 1 T
AUSTRALIA
1973
Douglas
Victoria Vi2
22 ?
Bald Hill
Phyllotheca? sp.
1
pl 14(3)
-
1
j Ti
1979
Holmes & Ash
Lome B T NE2
9 ! Camden Head
Camden Head (1583)
circular ribbed object
1
pl 3(8)
-
1
44 1
1982
Holmes
Dubbo | Syl
18 (unnamed)
Benolong (Ugothery)
Peltaspermum
1
pl 8(E,F)
-
1 1
1
tli
1986
White (specimen repeated from Holmes 1982)
ph on p 153(232)
-
*
" 1
assemblages
(taphocoenosis)
Genera
Species
Intactness
Molteno Cycles
Lepidopteris
1
£
3
E
<D CTJ
8* 5
2 0)
Q) C
Q. st
9 \d
P. monodiscum
P. tridiscum
P. turbanatum
P. quindiscum
Intact strobili
Fragmentary strobili
Isolated discs
Cal 111
Dic/Sph
5
:
“I “
1 |
I
“1
Bir 211
Sph 2spp
1
"l
1: -
-|
1 -
1
“l
1
5
” 111
70
67 -
57
- 10 -
8 4
55
Boe 111
Lep sto
90 - -
-1 -1 -
4/5
Tel 111
Hei elo
1
14 -
14.
-i -
4: 5
5
Ela 111
Die odo
51
3' -
3!
3' -
-
Kra 111
Die odo
10
1 -
1
1 1
1
-
3
Lut 311
Hei elo
19
7 -
7;
-j -
2
5
Tin 121
Sph 2spp
1
_! _
-
_i
Wal 111
Die odo
2: -1 -
-! -1 -1 -
-1 -! -
2/3
Kon 223
Die odo
4
-1 “
-i -1 -
" 222
9
24 2
24 -
8 5
11
” 111
24:
2, -
-j
2: - -
2; -
-
2f
Pen 321
Dic/Ris
9
V -
J
1 - -
1 -
-
" 421
Die odo
1
-1 -
-1 -1 .
-1
Kle 111
Hei/Dic
1
5 -
5!
-1 -
1 2
2
Kap 111
Dic/Ris
2!
I! -
1
r t
-j 1
-
2e
Mak 111
Die odo
7
1
1
Maz 111
Die era
2
2c
" 211
Hei/Dic
1
9 18
9.
-1 -
-1 -
9
Hla 211
Die 3spp
1
-1 "
-!
" 212
Die 3spp
1
1
"i "1
“I
” 213
Die elo
1
1 1
2b
Umk 111
Die 2spp
1
17 7
- 17
3 1
13
San 111
Die era
1
2 1
-1
.1 .1 2
-1 1
1
Mat 111
Die dub
1
4 -
-! -I 4
-1 1
3
Lit 111
Dic/Hei
1
11 4
10
- - d
4 -
7
2a
Aas 411
Dic/Sph
7
88 -
5
83 -
22 16
50
” 511
Die elo
10
-1 _
1 I
_(
Ask 111
Equ sp
20
-1 -
.1 .1 .
-1
Bam 111
Die dub
l!
-! -
-! -
Total TCs
30
17 5
10
3 3 4
11 10
12
Total indivs
% 257 32
112
27 94 24
57 38
162
assemblages
(taphocoenosis)
Genera
Species
Intactness
Habitat
Lepidopteris
E
3
i
8.
<0
£
9;
•2
c/5
Q)
C
d
P. monodiscum
P. tridiscum
P. turbanatum
P. quindiscum
Intact strobili
Fragmentary strobili
Isolated discs
Umk 111
Die 2spp
1
17!
7
-
1
17
3: 1!
13
D1
Lit 111
Dic/Hei
1
11
4
10
_ 1 J
1
4 -!
7
Hla 213
Die elo
i:
->
- -
” 212
Die 3spp
1
-
.1 -1
” 211
Die 3spp
1
-i -
-!
Mat 111
Die dub
1
4;
-
-
-! -
4
-! 1:
3
n?
Maz 111
Die era
2
-J
” 211
Hei/Dic
1
9
18
9
_ : _l
-
-;
9
Kap 111
Dic/Ris
2'
1
-
1
-1 -
-
1
-
San 111
Die era
1
2
1
-
-! -
2
-! 1
1
Kon 222
Die odo
9
24
2
-
24 -
-
8 5
11
" 223
4
-]
- | -
1 1
" 111
24
2;
-
-
2 -
-
2; -:
-
Pen 321
Dic/Ris
9
It
-
-
1 -
-
1 1
-
” 421
Die odo
1
-!
-1 -
-1 “I
D3
Ela 111
”
5
3
-
3
-
3 -!
-
Kra 111
10
1
-
1
-
-
i! -!
-
Mak 111
7
-|
-
Bam 111
Die dub
1
-1
- 1 -
Cal 111
Dic/Sph
5
-!
-
Wal 111
Die odo
2
-1 -
Aas 411
Dic/Sph
7
88
-
5
-83
-
22; 16
50
” 511
Die elo
10
- | “
Bir 111
Sph 2spp
70
67
-
57
-1 10
-
8 4
55
” 211
”
-!
1
-
-
-! 1
-
-I -1
1
Tin 121
1
_]
-! -
Lut 311
Hei elo
19
7
-
7
-: -
-
2
5
Tel 111
1
14
-
14
-| -
-
4 5
5
H
Kle 111
Hei/Dic
1
5
-
5
.1 -
-
1 2
2
Ask 111
Equ sp
20
-1 -1 "I -
-! -1 -
E
Boe 111
Lep sto
90
?
Total TCs
30
17
5
10
3: 3
4
11 10
12
Total indivs
%
257 32
112: 27 94
24
57 38
162
Tab. 40. PeltaspermumILepidopteris, Molteno occur-
rence, emphasizing stratigraphy (cycles, members)
Tab. 41 . PeltaspermumILepidopteris, Molteno
occurrence, emphasizing ecozones (habitat)
Habitat codes: see tab. 47
Peltaspermum
PELTASPERMALES
g/tRELITZIA 15 (2003)
151
Intactness and preservation of cones
Degree of cone fragmentation
Of 257 Peltaspermum individuals in our Molteno collection, 57 con-
sist of more or less intact strobili, 38 are fragmentary strobili and the
remaining 162 isolated dehisced discs (Tabs 40, 41). Recorded as relative
percentages, the three categories rate as 23% intact, 15% fragmentary. 62%
dehisced discs. We have nowhere attempted, on site, to estimate the rela-
tive percentages of intact individuals as preserved, but since our sampling
bias has strongly favoured the more completely preserved material, the fig-
ure might be as low as 5 or 10%. This would be comparable, for instance,
to Teletnachits.
Habitat bias
A particularly high proportion of intact strobili occur in the Dicroidium
open woodland habitat, most notably in the case of P. tridiscum in the wide-
spread cherty beds of the Konings Kroon/Peninsula district (D3 in Tab. 41 ).
The most likely conclusion, perhaps, is that these plants grew relatively
near to the sites of deposition and were deposited under moderate flow con-
ditions. At the other end of the scale, though Lepidopteris foliage occurs
regularly at around 1% of the assemblage in the immature Dicroidium
riparian forest TCs, intact Peltaspermum strobili are absent in these sites
(D2 in Tab. 41).
In situ seeds
Following the pattern observed in Teletnachits, the great majority of
Peltaspermum strobili, whether intact or fragmentary, have lost their seeds.
Rare specimens with in situ seeds have been recovered from Kon 222, pi.
40( 1—4) and Umk 1 1 1 . pi. 44(2, 5).
Dispersed seeds
The typical elongated tetrahedral seeds with bifid tips occur common-
ly in several TCs, particularly Aas 411, pi. 41(6, 7), pi. 42(6-8). We have
not attempted to systematically tabulate their occurrence.
Species nomenclature in Peltaspermum
The uncertainties associated with species nomenclature must be
addressed in naming the four clearly defined species that we recognise in
our Molteno collections. The problem is that the holotypes nominated in
Thomas (1933, 1955), Harris (1937) and Townrow (1960) often do not
show the diagnostic features that are available to us in our far more exten-
sive and often better preserved collections.
Pteruchus edwardsii Thomas (1933), later described as Stachyopitys
edwardsii (Thomas) Townrow (1960), was based on a single intact stro-
bilus from Konings Kroon. It is probably a Peltaspermum. It is uncertain
from which of our TCs in the region the material of Thomas may have been
collected. The specimen apparently has simple megasporophylls bearing
single discs, not three as in our Konings Kroon palaeodemes. As the preser-
vation is insufficient to make comparisons with our material, we here
regard Thomas’s material as Peltaspermum sp. indet.
Lepidopteris natalenisis Thomas (1933) was instituted by Thomas for
a partial, intact inflorescence from his ‘Bumera Waterfall locality' (our
Umk 111). The collections made by Thomas are housed in the Natural
History Museum. London. That specimen was renamed Peltaspermum
thomasii by Harris (1937) who provided a diagnosis, but no illustrations.
The type and two other specimens show simple megasporophylls bearing
single discs. From Umk 1 11, we have collected only intact specimens with
compound megasporophylls or diagnostic dehisced discs falling in the
species P. quindiscum.
P. thomasii is retained as a distinct species. It is closest to P. monodis-
cum in that both bear single discs. The discs of P. monodiscum differ from
those of P. thomasii in the deeply dissected and more numerous lobes. For
P. thomasii , Townrow (1960) recorded two seeds per disc, but there is prob-
ably one seed per lobe as in P. rotula from Greenland and in our new
Molteno species.
Poort & Kemp (1990) created the new ‘natural genus’ (their term)
Meyenopteris for the megasporophyll Peltaspermum thomasii plus the leaf
Lepidopteris stormbergensis ( natalensis ). We see no advantage in using
their name at present. They also defined the megasporophyll as bearing
only two ovules and being bilaterally symmetrical, while our specimens are
clearly multi-ovulate and radially symmetrical.
Adaptive radiation (Molteno diversity)
Diversification in the Peltaspermum! Lepidopteris plant is most evi-
dently expressed in the female strobilus, with five species described from
the Molteno. This contrasts markedly with the male affiliate, Antevsia, with
only one species recognised. The diagnostic characters defining the species
are the number of peltate discs (1 to 5) per megasporophyll; the number,
shape, degree of division, ornamentation and attachment of the lobes; and
the prominence and shape of the apical cap.
At least five species can be differentiated, four from Dicroidium- dom-
inated habitats and one from a Sphenobaiera closed woodland habitat.
They are based on the following TCs/reference palaeodemes:
P. thomasii— Umk 111 Die 2 spp (Umkomaas Valley), 3 indivs
Dicroidium riparian forest (mature); Cycle 2b (Indwe Member)
P. monodiscum— Mot 111 Die odo (Morija) 50 indivs
Dicroidium open woodland (floodplain); Cycle 2c (Indwe Member)
P. tridiscum— Lit 111 Dic/Hei (Little Switzerland) 15 indivs
Dicroidium riparian forest (mature); Cycle 2a (Indwe Member)
P. turbanatum— Aas 411 Dic/Sph (Aasvoelberg), 60 indivs
Sphenobaiera closed woodland (floodplain lake); Cycle 1 (Bamboesberg Member)
P. quindiscum— Umk 111 Die 2 spp (Umkomaas Valley), >16 indivs
Dicroidium riparian forest (mature); Cycle 2b (Indwe Member)
Peltaspermum thomasii T.M.Harris 1937
Species concept
A Peltaspermum species with simple megasporophylls bearing undivid-
ed discs with ca 6 lobes.
Comment & comparison
This species was originally described by Thomas (1933) from Umk
111, later named by Harris (1937) and further described by Townrow
(1960).
P. thomasii is distinct from all other Peltaspermum species from the
Molteno (see generic text for further comment). We have found no further
specimens of this species in our particularly extensive collections from the
original site (Umk 111).
Peltaspermum thomasii (from Townrow, 1960)
V. xl
V3579Sb
> ' ;;
w
xl
%
Umk 111
Holotype
PELTASMPERMALES
Peltaspermum
152
^/tRELITZIA 15 (2003)
Peltaspermum monodiscum j.M.And. & H.M.And., sp. nov.
Holotype
Specimen : PRE/F/10543; pi. 37(1, 5, 6).
Assemblage (TC): Bir 1 1 1 Sph 2spp. Birds River.
Preservation: virtually complete strobilus, without counterpart, with in situ
seeds in a few discs; impression in thinly laminated, yellowish grey
shale with very good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 57 indivs (7 intact, 1 partial, 49 isolated discs), pi. 37.
Sister palaeodemes— 9 (best 3 listed)
Tel 111 Hei elo: 14 indivs (4 intact, 10 isolated discs).
Lit 111 Dic/Hei: 10 indivs (4 intact, 6 isolated discs).
Aas 411 Dic/Sph: 5 indivs (1 intact, 4 isolated discs), pi. 38(1^1).
Specific diagnosis
A Peltaspermum species with simple megasporophylls bearing single
deeply divided, linear-lobed discs without cap.
Specific characters
Megasporophyll: simple, bearing a single disc.
Disc: lobes 1 1 or 12, linear, deeply divided, longitudinally grooved; apical
cap not developed.
Etymology
monodiscum— with reference to the single disc per megasporophyll.
Comment & comparison
Represented by 10 palaeodemes in the Molteno, this is the most fre-
quently occurring Peltaspermum species in the formation. It is very dis-
tinctive and, except for P. thomasii , shows no overlap with palaeodemes of
other species. Apical caps (which appear clearly in the other three species)
have not been observed in P. monodiscum.
Peltaspermum tridiscum j.M.And. & H.M.And., sp. nov.
Holotype
Specimen: BP/2/4137a,b; pi. 39(1, 2, 8)
Assemblage (TC): Kon 222 Die odo, Konings Kroon
Preservation: virtually complete strobilus, part and counterpart, no seeds;
clearly preserved; 3D mould and cast in thinly laminated, medium grey
cherty shale with poor cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 24 indivs (8 intact, 5 partial, 1 1 isolated discs), pis 39, 40.
Sister palaeodemes — 2 (both listed)
Kon 1 1 1 Die odo: 2 indivs (2 intact).
Pen 321 Dic/Ris: 1 indivd intact).
Specific diagnosis
A Peltaspermum species with compound megasporophylls bearing 3
partially divided, roundly lobed discs with cylindrical cap.
Specific characters
Megasporophyll: compound, bearing 3 discs.
Disc: lobes 10 or II, partially divided, with rounded ends; apical cap cylin-
drical.
Etymology
tridiscum— with reference to the three discs per megasporophyll.
Comment & comparison
The species is confined to the extensive cherty horizon of the Konings
Kroon-Peninsula district, and it is the only member of the genus found in
these cherts. Though P. turbanatum also has megasporophylls generally
bearing three discs, the features of the discs are clearly distinctive. The api-
cal cap is rarely preserved and is best seen — positive and negative— in the
two specimens BP/2/4144a,b and BP/2/4140a,b, pi. 39(5). The strobilus
axis has a characteristic zig-zag not found in other species.
Peltaspermum
PELTASPRERMALES
c/tRELITZIA 15 (2003)
153
Peltaspennum turbanatum J.M.And. & h.m And., sp. nov.
Holotype
Specimen'. PRE/F 21440a,b; pi. 41(1, 2).
Assemblage (TC): Aas 41 1 Dic/Sph, Aasvoelberg.
Preservation', virtually complete strobilus, part and counterpart, no in situ
seeds; impression in thinly laminated, strongly baked, yellowish grey
shale with very good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 83 indivs (21 intact, 16 partial, 46 isolated discs), pis 41, 42.
Sister palaeodemes— 2 (best 1 listed)
Bir 111 Sph 2spp: 10 indivs (1 intact, 3 partial. 6 isolated discs), pi. 43(1-6).
Specific diagnosis
A Peltaspermum species with compound megasporophylls bearing 3
or 4 undivided, roundly lobed discs with large turban-like cap.
Specific characters
Megasporophyll: compound, bearing 3 or 4 discs.
Disc: lobes 10, short/stout, not divided or grooved, extending from dis-
tinctive concentric collar; apical cap pronounced, turban-like.
Etymology
turbanatum— with reference to the turban-like apical cap.
Comment & comparison
P. turbanatum is restricted to the lake deposits of Aasvoelberg and
Birds River. Though the single intact specimen from Bir 111 (tf. 2) clearly
shows at least one megasporophyll bearing four discs— and this may well
have been the norm— the features of the individual discs are very like those
of Aas 41 1 and the palaeodeme is therefore included in the same species.
At Aas 41 1 the isolated tetrahedral seeds [pis 41(6, 7), 42(5, 7, 8)] are
very common and occur on most slabs.
Peltaspermum quindiscum J.M.And. & h.m. And., sp
nov.
Holotype
Specimen: PRE/F/ 1 87 11 a, b; pi. 44(1, 2, 4—6).
Assemblage (TC): Umk 111 Die 2spp, Umkomaas Valley.
Preservation: virtually complete strobilus, part and counterpart, with in
situ seeds: compression in thinly laminated, carbonaceous (good cuti-
cle), moderately baked, dark grey shale with good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 17 indivs (3 intact, 1 partial, 13 isolated discs), pi. 44.
Sister palaeodemes— 3 (best 1 listed).
Lit 111 Dic/Hei: 1 indiv. (intact).
Specific diagnosis
A Peltaspermum species with compound megasporophylls bearing ca
5 undivided, truncate-lobed discs with short conical cap.
Specific characters
Megasporophyll: compound, bearing 5 discs.
Disc: lobes 10, not divided or grooved, truncate; apical cap conical.
Etymology
quindiscum— with reference to the five discs per megasporophyll.
Comment & comparison
P. quindiscum is confined to four palaeodemes representing riparian
forests in the lower sections of the Indwe Member. In view of the distinc-
tive nature of the discs, the material (mostly isolated discs) from San 111,
Mat 1 1 1 and Lit 111 has been placed in the same species as the reference
palaeodeme from Umk 111.
PELTASPERMALES
Peltaspermum
154
d^TRELITZI A 15 (2003)
Alltevsia T.M.Harris 1937
Type species
Antevsia zeilleri (Nath.) T.M.Harris 1937.
Scoresby Sound, E. Greenland; Rhaetic, Triassic.
Generic concept
A ginkgoopsid male strobilus of variously branched form, with linear,
planar, simple to branching microsporophylls bearing lateral, sessile clus-
ters of microsporangia.
Generic characters (Molteno Fm.)
Attachment', strobilus borne singly on a bulbous short shoot (which dehisces
as a unit).
Strobilus: simple, lax to compact, bushy, radially symmetrical, medium-
sized ( ca 80 X 70 mm); axis erect, tapering; microsporophylls fairly
numerous, irregularly helical.
Microsporophyll: simple to irregularly forked or branched, linear, planar,
pinnate; bracteoles absent; rhachis robust, with clear distinction into
blistered midrib and naked flange; fertile heads numerous, regularly
closely spaced, sessile, in opposite to subopposite pairs; microsporan-
gia fascicled to pinnulate, 3-10 per head.
Microsporangium: irregularly elliptic (2-3 mm long); ornamentation fine,
linear, sinuous, forking and converging.
Pollen: nonsaccate. monocolpate.
Eponymy
Antevsia— after Dr E. Antevs, a Swedish palaeobotanist who added greatly
to the understanding of this genus.
Global range: numerous species, Pangaea, Tr.-J.
Gondwana Triassic occurrence
SAf— Karoo Basin, 5 TCs (33 indivs).
Antevsia appears not to have been previously recorded from elsewhere
in the Gondwana Triassic (except incorrectly by Townrow 1960).
Molteno occurrence
Frequency (F): 5 TCs (of 100 sampled in the Molteno).
Diversity (D): 1 species.
Abundance (A): 32 individuals, very rare to extremely rare.
Maz 211 Die 3spp:
18 indivs
in 85 man-hrs cleaving (2 per
1 man-day) very rare
Kon 222 Die odo:
1 ”
” 40
” (1 ”
2 ” )
San 1 1 1 Die era:
2 ”
” 30
” (1 ”
1 ” )
Umk 111 Die 2spp:
7 ”
” 400
” (1 ”
6 ” ) extr. rare
Lit 111 Dic/Hei:
4 ”
” 550
” (1 ”
14 ” )
Antevsia is far less common overall and less frequent than the female
affiliate Peltaspermum.
Affiliated organs
Female strobilus: Peltaspermum— Grade 3 (Kin. reinf., Mut. occ.).
Foliage: Lepidopteris— Grade 3 (Kin. reinf., Mut. occ.. Mor. corr.).
Classification & comparison
Suprageneric classification (Peltaspermaceae/Peltaspermales)
See discussion under Peltaspermum (p. 148).
Intergeneric comparison (Gondwana Triassic)
Antevsia is close to the genera Stachyopitys and Pteruchus in that all
bear clusters of fairly similar microsporangia. Poorly preserved specimens
can easily be confused. However, Antevsia has sessile lateral clusters of
3-10 microsporangia; Pteruchus is differentiated by the clear laminate
head bearing numerous abaxial microsporangia; Stachyopitys has numer-
ous microsporangia radiating from a central receptacle.
Townrow (1960), in error, classified Fanerotheca exstars as Antevsia.
Earlier, we followed Townrow and thus ascribed two specimens [And. &
And. 1983, pi. 23(3, 4)] to Antevsia, which in the present study are reclas-
sified as Fanerotheca.
Comparisons beyond Gondwana Triassic
Laurasian Jurassic— The Middle Jurassic genus Caytonanthus
(Crane 1985), first described from the Yorkshire flora of England,
has clusters of microsporangia similar to Antevsia. They differ in
Caytonanthus bearing multiloculate sporangia and Antevsia simple
sporangia.
Laurasian Permian— The Upper Permian (Tatarina Flora,
Russian Platform) genus Permotheca, placed in the order Pelta-
spermales by Gomankov & Meyen (1986), has clusters of simple
microsporangia similar to Antevsia. The pollen is recorded as disac-
cate by Meyen (1987, fig. 78d).
Reconstructions
The R4 reconstruction of A. mazenodensis (tf. 1 above), with many
unbranched microsporophylls angling steeply upwards from the central
axis, is based on the holotype (Maz 211, PRE/F/4653). The only other rea-
sonably sized portion of a strobilus (Maz 211, PRE/F/1 1729), intact except
for the base, shows a far more complex branching pattern of the
microsporophylls. A fascicle of three sporangia (Umk 111, PRE/F/6773) is
illustrated in tf. 2.
The curious reconstruction of a Lepidopteris leaf with an Antevsia-\\ke
strobilus attached to the distal extremity of the midrib was illustrated in
Taylor & Taylor (1993, fig. 15.31). This was based on two specimens col-
lected by Zavada (pers. comm., 1994) from Umkomaas Valley. As we have
never observed anything similar, we await a published description of this
unique find.
Evidence for affiliation of organs
Compared to the female strobilus Peltaspermum, the male, Antevsia, is
rare in the Molteno and co-occurs with Lepidopteris in only five TCs (Tab.
42). The affiliation of Antevsia with the female fruit Peltaspermum and the
foliage genus Lepidopteris merits Grade 3 reliability. This is supported by
the Laurasian evidence given below.
Mutual occurrence
Antevsia is found only in Cycle-2 of the Molteno and almost exclu-
sively (four of five appearances) in TCs associated with Dicroidium ripar-
ian forests (Tabs 4CM-2). In these Cycle-2 TCs the correspondence of
occurrence between Antevsia and Peltaspermum is near perfect, while else-
where in the Molteno the correspondence is almost nil. This discrepancy
may be explained by postulating differing seasons of maturation for the
male and female fruit.
Morphological correspondence
The Antevsia specimens from Umk 111 (most notably PRE/F/6773,
tf. 2 above) show the same distinctive blistering of the midrib that is found
in Lepidopteris.
Laurasian evidence
Antevsia occurs with Lepidopteris at three Swedish Rhaetic localities
(Antevs 1914) and at three Greenland Rhaetic localities (Harris 1932a),
where Peltaspermum is also present. Both Antevs and Harris have demon-
strated the cuticular similarities of the two organs.
Antevsia
PELTASPERMALES
C^TRELITZIA 15 (2003)
155
assemblages
(taphocoenosis)
Call 111
Bir 211
” 111
Dic/Sph
Sph 2spp
Boe 111 Lep sto
Tel 111
Ela 111
Kra 111
Lut 311
Tin 121
Hei elo
Die odo
Hei elo
Sph 2spp
Wal 111 Die odo
Kon 223
„ 222
” 111
Pen 321
” 421
Kle 1 11_
Kap 111 "
Mak 1lV
Maz 111
”211
Hla 211 *
" 212
” 213
Umk 111
San 111
Mat 111
Lit 111
Dic/Ris
Die odo
Hei/Dic
Dic/Ris
Die odo
Die era
Hei/Dic
Die 3spp
Die elo
Die 2spp
Die era
Die dub
Dic/Hei
Aas 411
” 511
Ask 111
Bam 111
Dic/Sph
Die elo
Equ sp.
Die dub
Total JCs
Total indivs
Genera
o TO
70 67
90
I 14
5 3
10 1
19 7
I I -
30 17
%257
d
4/5
2/3
Tab. 42. Antevsia, Molteno occurrence
PRE/F/6773b
pis 47(1— 7),48(4)
Antevsia mazenodensis j.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/4653; pi. 45(1-6).
Assemblage (TC): Maz 211 Hei/Dic; Mazenod.
Preservation : almost complete strobilus, without counterpart; compression
in thinly laminated, carbonaceous (poor cuticle), medium grey shale
with moderate cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 18 indivs (2 intact, 14 partial, 2 isolated microsporangia), pis
45(1-6), 46(1-7).
Sister palaeodemes— 4 (best 1 listed)
Umk 111 Die 2spp: 7 indivs (1 intact), pis 47, 48.
Specific diagnosis
An Antevsia species with microsporophylls in opposite fertile heads
bearing 3-6 microsporangia.
Specific characters: as per genus.
Etymology
mazenodensis— with reference to the type locality.
Comment & comparison
From the relatively few Molteno specimens at hand, the Antevsia stro-
bilus appears to show a particularly irregular morphological organisation
(tfs 1-3 below). In only one specimen (Maz 211. PRE/F/4653, tf. 2 below)
is the more or less complete strobilus preserved. It is attached to a bulbous
short shoot similar to that seen in many Stachyopitys individuals.
This Molteno species differs from the Faurasian Antevsia zeilleri,
which bears two or three ultimate fertile heads and clusters of 6-10
microsporangia. While five distinct Molteno species are recognised in the
ovulate genus Peltaspermum, the present collections are insufficient to
attempt to differentiate species within the supposed male affiliate Antevsia.
1-10: degre
O: superlo
e squares j
^lities j
ff/4
1
2
3k
&
4 ■•-Lk.
5
6
b/"\.
7 ■- "'ir'?
tT
$9
Molt
;no Fm.
PELTASPERMALES
Antevsia
156
Ci/TRELITZIA 15 (2003)
Lepidopteris Schimp. 1869
Type species
Lepidopteris stuttgardiensis (Jaeger 1827) Schimp. 1869.
Near Stuttgart, Germany; Keuper, Triassic.
Generic concept
A ginkgoopsid leaf with robust blistered rhachis, obovate, pinnate to
bipinnate lamina and zwischenfiedem.
Generic characters (Molteno Fm.)
Attachment : unknown.
Leaf, pinnate to bipinnate, obovate, medium to large; petiole short stout,
rhachis robust with characteristic blisters; pinnae opposite to suboppo-
site, closely spaced, sometimes decurrent, entire to deeply pinnatisect
to pinnate, narrowly lanceolate; zwichenfiedern usually present; pin-
nules (in bipinnate forms) opposite to subopposite, closely spaced,
often conjoining and becoming coherent; venation pinnate, lateral
veins fine, simple or forking; in coherent forms, sometimes conjoining
with veins of adjacent pinnules.
Cuticle: see And. & And. (1989, p. 90); this vol., tfs 1^1 below.
Etymology
Lepidopteris — lepido (Gr.), scaly; pteris (Gr.), fern.
Classification & comparison
Intergeneric comparisons
Gondwana Triassic— Lepidopteris, as a bipinnate leaf with intercalary
pinnules (zwischenfiedem) along the rhachis, is unique amongst the
Gondwana ginkgoopsid leaf genera. Lepidopteris and the genus Scyto-
phyllum are end members of an intergrading range of leaves in which the
pinnules coalesce and become coherent. The choice of genus for leaves that
are partially coherent is subjective. The cuticular features compare well
with those of other ginkgoopsid foliage, and most closely resemble
Dicroidium. Lepidopteris cuticle is well-developed, with notable features
such as buttressed cell walls and the ring of subsidiary cells. Where mate-
rial is sparse or imperfectly preserved, Lepidopteris may be confused with
fern leaves.
Interspecific comparisons
Of the five Gondwana Triassic species of Lepidopteris recognised by
us (And. & And. 1989), only L. stormbergensis and L. africana occur in the
Molteno. The two forms are found together in 1 1 of the 30 Molteno TCs
yielding Lepidopteris, and in most of these cases they appear to constitute
an unbroken morphological range within a single palaeodeme. In the
remaining TCs, only one or the other of the species is encountered.
Global range: several spp.. Pangaea, U.P.-U.Tr.
Gondwana Triassic occurrence (after And. & And. 1989)
Frequency (F): 19 degree squares (of the 84 across Gondwana).
Ubiquity (U): 4 continents (of 5 comprising Gondwana).
Diversity (D): 5 foliage species.
Abundance (A): 1% (the norm in Molteno TCs).
Longevity (L); 21 myrs (Scythian to early Norian).
Colonisation success: FUDAL rating 19/4/5/1/21 = 50.
Intermediate success (Grade 3): Lepidopteris was the eighth most
prominent genus in the Gondwana Triassic; it was frequent, ubiquitous
and long-lived, but of relatively moderate diversity and abundance.
Endemism: of the five described Gondwana Triassic species, three (L.
africana, L. stormbergensis and L. madagascariensis) are widespread,
while the remaining two (L. brownii and L. langlohensis), as known,
are single-formation endemics.
Molteno occurrence
Frequency (F): 30 TCs (of 100 sampled in the Molteno).
Diversity CD): 2 species.
Abundance (A): monodominant (90%) in 1 TC; co-dominant (20%)
in 1 TC; common to abundant (5-10%) in 3 TCs; occasional ( 1%)
in 10 TCs; rare to very rare (<1%) in the other 15 TCs.
Habit: possibly woody, much-branched spreading shrub.
Preferred habitat: ubiquitous in Dicroidium riparian forest, less fre-
quent (5 of 10 TCs) in closed woodland of the lake margins.
Affiliated organs
Female strobilis: Peltaspermum— Grade 4 (Mut. occ., Mor. cor.. Cut. cor.).
Male strobilus: Antevsia-Gta.de 3 (Kin. reinf., Mut. occ.).
L epido pteris stormbergensis
Lit 111 PRE/F/5610
upper
lower
tfs 1-4
from And. & And , 1989
lower
Lepidopteris
PELTASPERMALES
^TRELITZIA 15 (2003)
157
N
L. africana
Lepidopteris
generic panorama showing
the 2 Molteno species
L. stormbergensis
, r ,6 Lit in
i \f\ PRE/F/5592
1-8 from
d & And.. 1989
it'
PRE/F/1897
GONDWANA TRIASSIC, GEOSTRAT DISTRIBUTION
1-10: degr
O: superlo
?e squares j
:alities C'
fls-
Pi
2
ir'"
3k
4 ' -U^
5
}-U 6
7
8 Qd f
9
ff 10
Molt
;no Fm.
SAm SAf Ind Ant Aus
Ch I NA I SA I Pa Lu Za U Ka WH| PI A* NZ| Ca | SA Ga Bo CM| NE Sy Vl Ta
(\
o: productive degree
squares
w?
Gondwana Triassic
PELTASPERMALES
Lepidopteris
c/trelitzi A 15 (2003)
\ ' * «%t-
PRE/F/10543 \
Holotype
PRE/F/15592
Birds River
(Bir 1 1 1 Sph 2spp)
PRE/F/10543
Peltaspermum monodiscum
pi. 37
PELTASPERMALES
r/TRELITZIA 15 (2003)
159
PRE/F/21443a
HBK' 5
PRE/F/21443b xl
Aasvoelberg
(Aas411 Dic/Sph)
P. monodisciim
Umkomaas Valley
(Umk 1 1 1 Die 2spp)
Holotype
j V23400 NHM London
P. thomasii
PELTASPERMALES
pi. 38
Peltaspermum spp,
160
d/TRELITZIA 15 (2003)
PRE/F/20313a
Konings Kroon Hoiotype
(Kon 222 Die odo)
?BP/2/4140b
Peltaspermum tridiscum
pi. 39
PELTASPERMALES
TRELITZIA 15 (2003)
161
PP.E/F
PRE/F/ 20000 b
PRE/F/20000b
PELTASPERMALES
pi. 40
Peltaspermum tridiscum
Konings Kroon
(Kon 222 Die odo)
PRE/F/20000b
PRE/F/20000a
162
d^TRELITZIA 15 (2003)
■fejs*
PRE/F/20665
PRE/F/2 1440a
PRE/F/20668a
PRE/F/20659
6 ’ a ' ! 1t§t
1X10 PRE/F/12802 XlO PRE/F/20668b
PRE/F/21440a
Holotype
Aasvoelberg
(Aas411 Dic/Sph)
PRE/F/20683
pi. 41
Peltaspermum turbanatum
PELTASPERMALES
'-■'/tRELITZIA 15 (2003)
163
PELTASPERMALES
pi. 42
Peltaspermum turbanatum
164
Peltasperm um turbanatum
pi. 43
PELTASPERMALES
rV TRELITZIA 15 (2003)
PRE/F/10017b
BP/2/B174
Birds River
(Bir 1 1 1 Sph 2spp)
PRE/F/10017b
c/tRELITZI A 15 (2003)
165
m§4
tX&H
PRE/F/1871 lb •(
Holotype
PRE/F/1871 lb
Umkomaas Valley
(Umk 1 1 1 Die 2spp)
^PRE/F/814b
) tl •• r • . • '
PRE/F/1871 lb',
PRE/F/1871 lb
PRE/F/1871 lb
PELTASPERMALES
pi. 44
Peltaspermum quindiscum
166
^TRELITZIA 15 (2003)
all
PRE/F/4653
Holotype
Mazenod
(Maz 211 Hei/Dic)
Antevsia mazenodensis
pi. 45
PELTASPERMALES
PELTASPERMALES
pi. 46
Antevsia mazenodensis
168
d/TRELITZIA 15 (2003)
Antevsia mazenodensis pi. 47 PELTASPERMALES
169
<if'. TRELITZIA 15 (2003)
PELTASPERMALES pi. 48 Antevsia mazenodensis
170
' '/t R 1. 1. 1 T Z I A 15 (2003)
Scytophyllum J.B.Bornemann 1856
Type species
Scytophyllum bergeri J.B.Bornemann 1856.
Miilhausen, Germany; ?Keuper. Triassic.
Generic concept
A ginkgoopsid leaf with robust blistered rhachis, lanceolate, pinnate -
pinnatisect lamina and zwischenfiedern coherent with pinnae.
Generic characters (Molteno Fm.)
Attachment', unknown.
Leaf, pinnate-pinnatisect, lanceolate, medium-sized; petiole unknown;
rhachis robust with characteristic blisters; pinnae opposite to suboppo-
site, short, broadly tapering, apex rounded, with coherent pinna-like
zwischenfiedern; veins lepidopteroid, midrib tapering, secondary veins
fine and give rise to simple or forking tertiary veins which sometimes
conjoin.
Cuticles: see text adjacent.
Etymology
Scytophyllum— skytos (Gr.), leathery; phyllom (Gr.), leaved.
Global range: several spp., Pangaea, L.Tr.-L.J.
Gondwana Triassic occurrence
Frequency (F): 1 degree square (of the 84 across Gondwana).
Ubiquity (U): 1 continent (of 5 comprising Gondwana).
Diversity (D): 1 foliage species.
Abundance (A); <1% (as recorded for the Molteno).
Longevity (L): 1 myrs (Lower Camian).
Colonisation success: FUDAL rating 1/1/1/— /I = 4.
Minimum success (Grade 1): Scytophyllum, as understood here, was
one of the three least prominent genera in the Gondwana Triassic.
Endemism: the Molteno species is a single-assemblage endemic.
Molteno occurrence
Frequency (F): 1 TC (of 100 sampled in the Molteno).
Diversity (D): 1 species.
Abundance (A): 1 indiv., extremely rare.
Habit: possibly a shrub.
Preferred habitat: Dicroidium riparian forest.
Affiliated organs: unknown for Molteno.
Classification & comparison
Scytophyllum often forms an integrating series with Lepidopteris as
pointed out by Dobruskina (1975, p. 536), Holmes (1982, p. 22) and And.
& And (1989, pp. 65, 88). The species described here from Umk 111 has
the completely coherent form as is diagnostic for Scytophyllum.
Vittaephyllum Dobruskina 1975 (U. Permian to U. Triassic of the
USSR), from the general appearance of the lamina, is evidently related to
Lepidopteris, but in its forking frond it is reminiscent of Dicroidium.
Krassilov (1991, 1995, 1997) records some interesting ideas on the
question of the origin of angiosperm leaves from the coalescing of adjacent
veins as in Scytophyllum leaves.
Scytophyllum austroafricanum j.M.And. & H.M.And.,
sp. nov.
Holotype
Specimen: PRE/F/399; pi. 49(1-3).
Assemblage (TC): Umk 111 Die 2spp, Umkomaas Valley.
Preservation: nearly complete leaf, without counterpart; compression in
thinly laminated, carbonaceous (good cuticle), moderately baked, dark
grey shale with good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 1 individual.
Sister palaeodemes: nil.
Specific diagnosis
A Scytophyllum species with a coherent lamina of pinna lobes and
zwischenfiedern.
Specific characters
Leaf, length unknown, at least 140 mm long; petiole unknown, rhachis ca
3 mm wide; pinna lobes vary from small (7x7 mm) at base to larger
(20 x 12 mm) in central area, opposite to alternate, entire to sinuate,
with characteristic small triangular zwischenfiedern between and coa-
lescing with the larger pinna lobes; venation complex, with tertiary
veins that may coalesce when lying in close proximity to each other.
Etymology
austroafi'icanum— with reference to the South African origin of the species.
Comment & comparison
This single specimen, with coherent pinnae and complex venation, is
placed in the genus Scytophyllum. It is similar in part to S. vulgare from the
Late Triassic of the Eastern Urals (Dobruskina 1969; Krassilov 1997),
which is polymorphic. The Molteno species is different from partially
coherent forms of Lepidopteris africana in which the pinnae are usually
two to four times longer than wide. The two South American (Argentina)
records of Scytophyllum, S. neuburgianum and S. bonettiae (Zamuner &
Artabe 1990; Zamuner et al. 1999), are here regarded as more likely to
belong to the genus Lepidopteris.
Cuticles
Potential sample: Umk 111,1 indiv.
Macerated (this work): 1 indiv.
Presen’ation grade: Grade 3, tiny pieces.
Diagnostic characters: not available.
Comment: The reason for our preparing cuticle samples was to assess the
similarity between this single individual of S. austroafricanum and
Lepidopteris (both included in the Peltaspermaceae) on the one hand,
and the foliage genus Kurtziana (in the adjacent order) on the other.
The tiny fragments of Grade 3 cuticle were insufficient for compar-
isons.
Significance: of no aid in classification and affiliation.
Scytophyllum
PELTASPERMALES
171
dv^TRELITZIA 15 (2003)
PRE/F/399
Holotype
Umkomaas Valley
(Umk 1 1 1 Die 2spp)
PELTASPERMALES
pi. 49
Scytophyllum austroafricanum
172
d^TRELITZIA 15 (2003)
GINKGOOPSIDA S.V.Meyen 1984
MATATIELLALES J.M.And. & H.M.And., ord. nov.
MATATIELLACEAE J.M.And. & H.M.And., fam. nov.
Matatiella J.M.And. & H.M.And., gen. nov.
Type species
Matatiella rosetta J.M.And. & H.M.And., sp. nov.
Matatiele, Karoo Basin, S. Africa; Camian, Triassic.
Generic diagnosis
A ginkgoopsid strobilus of linear-cylindrical shape with megasporo-
phylls consisting of single, reflexed, variously lobed palmate heads.
Generic characters
Strobilus: simple, racemose, compact, radially symmetrical, of medium size
( ca 90 mm long); axis relatively stout, erect; megasporophylls numerous,
helically arranged.
Megasporophyll: simple, pedunculate; ovuliferous heads palmate (10 X 8
mm), recurved, 4-6-lobed; ovules/seeds (?)adaxial, enclosed by thin
protective membrane, 1 per lobe.
Ovule : oval (2 X 1.5 mm), unwinged.
Etymology
Matatiella— after the type locality Matatiele.
Global range: 5 spp., Gondwana, Tr. (LAD-CRN).
First: Matatiella sp. indet. (Retallack 1981b); Long Gully Fm., Benmore Dam
region. New Zealand.
Last: the 4 Molteno species described here.
Gondwana Triassic occurrence
SAf— Karoo Basin. 4 TCs (17 indivs).
Aus— New Zealand, 3 TCs (24 indivs).
Molteno occurrence
Frequency (F): 4 TCs (of 100 sampled in the Molteno).
Diversity ( D): 4 species.
Abundance (A); 17 individuals total, very rare to vanishingly rare.
Pen 41 1 Hei elo: 7 indivs. in 70 man-hours (1 per 1 man-day) very rare
Mat 111 Die dub: 6 ” ” 65 ” (1 ” 1 ” )
Kan 111 Ast spA: 3 ” ” 30 ” (1 ”1 ” )
Aas 4U Dic/Sph: 1 ” ” 512 ” (1 ” 51 ” ) vanish, rare
Matatiella is an infrequent and very rare element in the Molteno.
Notably, it is represented by a distinct species in each palaeodeme.
Affiliated organs
Male strobilus: unknown.
Foliage: Kurtziana— Grade 2 (Mut. occ.).
Classification & comparison
Suprageneric classification (Matatiellaceae/Matatiellales)
Of all known global Triassic megasporangiate strobili, Peltaspermum is
the only genus with some overall similarity to Matatiella. It might seem rea-
sonable to relate the multilobed, peltate, ovuliferous disc of Peltaspermum
with the lobed, palmate head of Matatiella. However, as the megasporophylls
of Peltaspermum are radially symmetrical and those of Matatiella bilateral-
ly symmetrical, their separation at order level appears to be justified.
Matatiella is placed in the new family Matatiellaceae and order Matatiellales
within the class Ginkgoopsida.
Intergeneric classification (Gondwana Triassic)
Considering the full set of seven Molteno ginkgoopsid megasporan-
giate genera— Peltaspermum, Matatiella , Avatia. Hamshawvia ,
Umkomasia , Kannaskoppia and Cetifructus— Peltaspermum, as noted
above, is the only strobilus at all like Hamshawvia. When individual
megasporophylls are viewed alone, then Avatia, with its palmate heads,
comes closest. These differ most evidently in bearing readily dehisced
winged seeds.
Reconstructions
The full R4 reconstruction is based on the two almost complete Mat
111 M. rosetta strobili shown on pis 50, 5 1 . As no clear tip is preserved, the
total length of the strobilus is uncertain. An apparent base is preserved in
PRE/F/10 193b— suggested by the absence of megasporophylls along the
axis (which, however, dips into the matrix). Possible tips are preserved in
the specimens from Kan 111 and Pen 411, but there are no clear bases.
In most specimens it is difficult to decide whether the ovules are
attached on the abaxial or adaxial side of the megasporophyll. The strobilus
reconstruction is drawn with the ovules in an adaxial position as that is how
they appear to be in the material from the type locality Mat 111. However, at
Kannaskop (Kan 111), a strobilus, PRE/F/ 13503a, is preserved showing a
cross section of a megasporophyll head, seemingly attached to the main axis
and with ovules apparently in an abaxial position— pi. 52(3); p. 174, tf. 4b.
Matatiella
MATATIELLALES
d/TRELITZIA 15 (2003)
173
Evidence for affiliation of organs
Mutual occurrence
We have given Matatiella a Grade 2 affiliation with Kurtziana based on
mutual occurrence in four Molteno TCs (Tab. 44). In these four TCs,
Kurtziana is always below 1% of the total flora although one would expect
higher numbers if Matatiella was indeed its affiliate. In the two TCs where
Kurtziana is quite common (Lut 511, 20% and Kon 111, 5%) no Matatiella
has been found. Our affiliation rating is supported by the New Zealand occur-
rences.
New Zealand
In New Zealand, Matatiella cournanei (formerly Peltaspermum cour-
nanei) has been affiliated with Pachydermophyllum praecordillera at three
TCs (Retallack 1981b, 1983; Pole & Raine 1994). Pachydermophyllum is
a northern hemisphere genus based on Yorkshire Jurassic fronds. The
Gondwana material requires revision. We consider it may be allied to, if not
the same as Kurtziana.
■Leyensis
Gondwana Permian glossopterids
The female glossopterid fruit, Righya and
Ligettoma from the Upper Permian of South
Africa (And & And., 1985, p. 127), show
definite similarities with Matatiella.
It is largely m the overall structure of
their strobili that they differ.
L. lidgettonoides
tfs 1-^t from And. &And., 1985
EstcourtFm., Karoo Basm, South Africa
L. africana
Adaptive radiation (Molteno diversity)
On the basis of the limited collection of Matatiella available— a total
of 17 individuals— a different species appears to characterise each of the
four TCs/reference palaeodemes. The affiliated leaf genus Kurtziana shows
much greater diversity, with some 16 species occurring. The four species
are distinguished by features of the megasporophylls, most notably the
number and nature of the ovuliferous lobes.
Each species represents a different habitat with a distinctive assem-
blage of plants, and derives from a different stratigraphical level.
M. rosetta— Mat 111 Die dub (Matatiele); 6 indivs
Dicroidium riparian forest (immature); Cycle 2b (Indwe Member)
M. hemirosetta— Kan 111 Die dub (Kannaskop); 3 indivs
fem IKannaskoppia meadow; Cycle 3 (Mayaputi Member)
M. sessilis— Pen 411 Hei elo (Peninsula); 7 indivs
Heidiphyllum thicket; Cycle 2f (Indwe Member)
M. reducta— Aas411 Dic/Sph (Aasvoelberg); 1 indiv.
Sphenobaiera closed woodland; Cycle 1 (Bamboesberg Member)
Gondwana Triassic occurrence (elaborated)
The specimens described by Pole & Raine (1994) from New Zealand
as Peltaspermum cournanei can now be placed in the new genus Matatiella
as M. cournanei (Pole & Raine) J.M.And. & H.M.And., comb. nov. They
show no distal extension to the ovuliferous lobe and are closest to M.
reducta described below.
Tab. 43.
MATATIELLA HYPODIGM, Gondwana Triassic occurrence
Species
Molteno
Other
in S v>
O ® <u
C -C <n
IIS
M. red.
M. cou.
sp. indet
AUTHOR
SUBREGION
FORMATION
LOCALITY
NAME
ILLUSTRATION
New Zealand
1
1
1
1 I
1 1
\
1981b Retallack
1983 1
Benmore dam
NZ4
21 \ Long Gully Farm
” j Bl. Jacks Congl.
Long Gully
nr. Benmore Dam
(?) Peltaspermum sp.
Peltaspermum
1 f 3(B)
3 f 6(F-H), f 10(G-I)
■ 1 " I
i
3? -
1994 Pole & Raine
Invercargill
?
? 1 ?
Pollack Road *
Antevsia
1 f 5(A), 6(D)
- 1 - 1 -
-
1? -
”
” i ”
”
P. cournanei
20 f 5(B-G), 7(A-G)
- : - I -
-
20 -
* all from same horizon, F46/F067
I
1
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
MATATIELLALES
Matatiella
174
^/tRELITZIA 15 (2003)
Matatiella VOSCtta J.M.And. & H.M.And., sp. nov.
Holotype
Specimen-. PRE/F/10193a.b; pis 50(1, 2, 5-7), 51(1-3).
Assemblage (TC): Mat 111 Die dub, Matatiele.
Preservation: central portion of cone, part and counterpart; impression, in
thickly laminated, olive-grey shale with moderate cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 6 indivs (2 intact, 3 partial, 1 isolated), pis 50(1-7), 51(1-5).
Sister palaeodemes— nil.
Specific diagnosis
A Matatiella species bearing nregasporophylls with 6 deeply divided
ovuliferous lobes forming a full rosette.
Specific characters
Megasporophyll: ovuliferous heads 6-lobed, forming a full rosette; lobes
deeply divided, distinctly narrowed towards base, extending well dis-
tal of the ovule and tapering to a truncate apex.
Etymology
rosetta— referring to the rosette form of the megasporophyll.
Comments & comparison
M. rosetta differs from other Matatiella species in the lobes which are
deeply divided to well beyond the ovules and almost to the centre of the
megasporophyll.
The high proportion of articulated and reasonably intact specimens of
this and other fruit taxa from Matatiele (Mat 111) suggests that it was an
autochthonous to near-autochthonous deposit.
Matatiella hemirosetta J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/13503a,b; pi. 52(1, 3, 7).
Assemblage (TC): Kan 111 Ast spA, Kannaskop.
Preser\’ation: central portion of cone, part and counterpart; impression, in
thick-bedded, moderately baked, greenish grey, silty mudstone with
very poor cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 3 indivs (1 intact, 2 partial), pi. 52(1-7).
Sister palaeodemes— nil.
Specific diagnosis
A Matatiella species bearing megasporophylls with 6 moderately
divided ovuliferous lobes forming a half rosette.
Specific characters
Megasporophyll: ovuliferous heads 6-lobed, forming a half rosette; lobes
moderately divided, extending well distal of the ovule to a broad trun-
cate apex.
Etymology
hemirosetta— referring to the half-rosette form of the megasporophyll.
Comments & comparison
M. hemirosetta differs from other Matatiella species in the semicircu-
lar shape and less deeply divided lobes of the megasporophyll.
The three articulated individuals of M. hemirosetta and the uniquely
intact preservation of other fruit and foliage taxa from Kannaskop (Kan 111)
suggest that this too (see text for M. rosetta) was an autochthonous deposit.
Matatiella
MATATIELLALES
c/trELITZIA 15 (2003)
175
Matatiella sessilis J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/1 7942a,b; pi. 53(5-7).
Assemblage (TC): Pen 41 1 Hei elo. Peninsula.
Preservation : a single megasporophyll head, part and counterpart; impres-
sion, in thickly laminated, greenish grey shale with moderate cleavage.
Reference palaeodeme
Assemblage (TC); as for holotype.
Specimens: 7 indivs (1 intact, 3 partial, 3 isolated), pi. 53(1-9).
Sister palaeodemes— nil.
Specific diagnosis
A Matatiella species bearing megasporophylls with 6 or 4 (2 aborted),
shallowly divided ovuliferous lobes.
Specific characters
Megasporophyll: ovuliferous heads 6- or4-lobed (sometimes with 2 ovules
aborted), forming a full rosette; lobes shallowly divided, extending
slightly beyond the ovule to irregularly rounded apex.
Etymology
sessilis— referring to the sessile megasporophyll lobes.
Comments & comparison
M. sessilis differs from other Matatiella species in the very shallowly
divided megasporophyll lobes which barely extend beyond the ovule. A
further characteristic, apparently unique to this species and none too certain
in view of the limited sample, is the tendency for some of the ovules to
abort (tf. 1 adjacent).
Matatiella reducta J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/20536; pi. 53(10, 11).
Assemblage (TC): Aas411 Dic/Sph, Aasvoelberg.
Preservation: a single megasporophyll head, without counterpart; impres-
sion, in thinly laminated, strongly baked, yellowish grey shale with
very good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 1 indiv. (isolated), pi. 53(10, 11).
Sister palaeodemes— nil.
Specific diagnosis
A Matatiella species bearing megasporophylls with 4 deeply divided
ovuliferous lobes not extending beyond the ovule.
Specific characters
Megasporophyll: ovuliferous heads 4-lobed, forming a full rosette; lobes
deeply divided, not extending beyond ovules.
Etymology
reducta— referring to the reduced megasporophyll lobes.
Comments & comparison
This species, based on a single specimen, is not as secure as the previ-
ous three described from the Molteno. It differs from those species in hav-
ing four megasporophyll lobes which do not extend beyond the ovules.
M. reducta is partly reminiscent of M. cournanei , originally described
as a Peltaspermum by Pole & Raine (1994) from New Zealand. That
species differs in having six ovulate lobes and is alike in that these show no
distal extensions beyond the ovules.
MATATIELLALES
Matatiella
176
d?TRELITZIA 15 (2003)
Kurtziana Freng. 1942
Type species
Kurtziana cacheutensis Freng. 1942.
Cacheuta, Argentina; Camian, Triassic.
Generic concept
A ginkgoopsid leaf with elliptical pinnate lamina and pinnules marked-
ly contracted above and decurrent below.
Generic characters (Molteno Fm.)
Attachment', unknown.
Leaf, pinnate, elliptical, medium-sized; petiole distinct, of moderate
length; pinnae generally closely spaced, opposite to subopposite, ovate
to broadly linear, entire, apex obtuse; base contracted above and vari-
ously decurrent below; veins simple to twice forked, midrib distinct to
apex.
Cuticle', see text adjacent.
Eponymy
Kurtziana— for F. Kurtz, an eminent South American palaeobotanist.
Global range: ca 20 spp., Gondwana, Tr.-J. (LAD-HET).
First: Kurtziana sp. (Walkom 1928); Esk Beds, Wivenhoe, Australia.
Last: Kurtziana brandmayri (Artabe et al. 1991); Nestares Fm., Alicura,
Neuquen Province, Argentina.
Gondwana Triassic occurrence (after And. & And. 1989)
Frequency (F): 6 degree squares (of the 84 across Gondwana).
Ubiquity (U): 2 continents (of 5 comprising Gondwana).
Diversity (D): ca 20 foliage species.
Abundance (A): <1% (the norm in Molteno TCs).
Longevity (L): 2 myrs (Lower Ladinian to Liassic).
Colonisation success: FUDAL rating 6/2/20/-/2 = 30.
Intermediate success (Grade 3): Kurtziana was the 12th most promi-
nent genus in the Gondwana Triassic; it was diverse, but of moderate
frequency and abundance.
Endemism: the 20 species are mainly single-assemblage endemics.
Molteno occurrence
Frequency (F): 13 TCs (of 100 sampled in the Molteno).
Diversity (D): 16 species.
Abundance (A): co-dominant (20%) in 1 TC; common (5%) in 1 TC;
rare to very rare (<1%) in other 1 1 TCs.
Habit: possibly a small spreading tree.
Preferred habitat: on the periphery of Heidiphyllum thicket.
Affiliated organs
Female strobilus: Matatiella , Grade 2 (Mut. occ.).
Male strobilus: not known.
Classification & comparison
Intergeneric comparison
In frond morphology and venation pattern, Kurtziana differs from all
other Gondwana ginkgoopsid leaf genera. Certain northern hemisphere
genera have previously been used for Gondwana leaves, some of which,
e.g. Pachypteris and Pachydermophyllum, may be better placed in
Kurtziana.
Kurtziana was placed in the Cycadales by Artabe & Stevenson ( 1999).
This was based on K. brandmayri (leaf and cuticle) from the Jurassic of
Neuquen Province, Argentina, as described by Artabe et al. (1991).
However, we feel there is no clear case for cycadalean affiliation based
only on cuticle which does not show clearly preserved stomata and sub-
sidiary cells.
In view of the possible affiliation (Grade 2) with Matatiella megas-
porophylls, we place Kurtziana in the Ginkgoopsida.
Interspecific comparison
Kurtziana has not been studied systematically on a Gondwana-wide
basis and the specimens included here may not all belong to the same nat-
ural genus. Cuticular studies should prove useful in indicating whether
Kurtziana. as conceived here, is indeed a natural genus or a form-genus
with many unrelated species. While we have included the type species,
K. cacheutensis, in the distribution map, the following two possible records
from Australia have not been plotted:
Thinnfeldia eskensis [Walkom 1928, pi. 28(1)]; Esk beds, Wivenhoe, Queens-
land.
Dicroidium eskense [Flint & Gould 1975, pi. 2(3)]; Basin Creek Fm., Nym-
boida, N.S.W.
From our Molteno collections we have provisionally recognised 16
Kurtziana species (Tab. 44). They remain undescribed and unnamed. Five
are illustrated opposite.
Cuticles
Potential sample: Umk 111, 18 indivs.
Macerated (this work): 6 indivs.
Preservation grade: Grade 3.
Diagnostic characters: marginally present.
Comment: some cellular structure available from 5 indivs, but not yet studied.
Significance: of no obvious aid in classification or affiliation.
assemblages
(taphocoenosis)
Kurtziana |
Species
-2
2
i
?
Species
Intactness
K. spA (Mat 111)
” sp B (Boe 112)
” sp C (Boe 112)
” sp D (Kon 111)
” sp E (Vin 111)
” sp F (Umk 111)
” each. (Umk 111)
” sp G (Hla 213)
” sp H (Mat 111)
” spl (Umk 111)
’’ sp J (Mat 111)
5
c >
0
CL
s
< n
” spL (Lut 511)
” sp M (Kan 111)
” sp N (Lut 311)
” sp O (Umk 111)
M. rosetta
M. hemirosetta
M. sessilis
M. reducta
Intact strobili
Fragm. strobili
Isolated heads
Boe 112
Die cor
3
-! 2; -] -|
1 1 1
1 1 1
“1 "1 "1
I 1 1
1 1 1
"1 "1 ”1
i
1
1 1 I
1 1 1
-1 "I “1
1 1 1
“1 - 1 " 1 -
1 1
- 1 - | -
Kan 111
Ast spA
2
-| - -| -| -|
-! -! -!
1
"1
J 9 - -
.. 1 ]
3
1 1 2 ; -
Vin 111
Die odo
7
~ _ 1 .1 .1 _l7
_i .1 J
J
.1 .1 .1 .
1 1
Kra 111 ” "
4
.1 .1 -1 .1 -1
-l4 -1
-i _i j
j
_i _i _i
.1 .1 - 1 -
_ 1 _ 1 _
Lut 511
Hei elo
20
-1 -| “I -| -1
-1 -1 -1
-1
20 - - -
-! - ! - 1 -
- 1 - 1 -
" 311 ” ”
5
- i - “ ) " ; 4
■ 1 -1 -!
-| -l 1
“i
-; -I 1! ■-
-1 - 1 - 1 -
- 1 - 1 -
Kon 111 ” ”
5
tz
l l 1
1
- 1 - | -
Pen 411
Hei elo
50
-|
_ 1 _ l j
50
7
-! - ; 7 : -
-; 4 3
Hla 213
Die elo
1
- 1 -1 -1 -1 -1
-1 -1 11
J
-1 -I .1
-1 - 1 - 1 -
.1 _ 1 _
Umk 111
Die 2spp
18
-! -! -! 2! -!
1 2i -!
-! 4 2'
-1
-1 -1 -1 6
-
-1 - ! - 1 -
_ 1 . ! .
San 111
Die era
1
-1 -1 -1 -1 -1
- 1 -
"
-1 - 1 - 1 -
- 1 - 1 -
Mat 111
Die dub
3
9 1 1 1 i T
J
-! -i - -
6
6 - 1 ! I
2 ; 3 ; 1
Aas 411
Dic/Sph
15
1 1 1 J _ i J -1
1
-! - 1 - 1 1
- ; - 1 1
Totals TCs
13
l! 1| L 2| 2\
1; 3! i;
1 1! 2;
3
1! 1: 1; 1
4
1; 1 ; 1 ; 1
2 ! 3 ; 3
Total indivs
%
2 2 1 % 11
1| 7! i;
I! 4 3|
65j
%! 2! 1 ! 6
17
6; 3; 7 ; 1
1 1
3] 9 ; 5
! 1
Tab. 44. Matatiella! Kurtziana, Molteno occurrence
Kurtziana
MATATIELLALES
TRELITZIA 15 (2003)
177
MATATIELLALES
Kurtziana
^TRELITZIA 15 (2003)
PRE/F/1 0193b
Matatiele
(Mat 1 1 1 Die dub)
Holotype
PRE/F/10193a
• PRE/F/10193a
X20
Matatiella rosetta
pi. 50
MATATIELLALES
179
^TRELITZIA 15 (2003)
PRE/F/10193a
PRE/F/10193b
Matatiele
(Mat 1 1 1 Die dub)
PRE/F/9257
PRE/F/10 193a'
pi. 51
MATATIELLALES
Matatiella rosetta
180
d?TRELITZIA 15 (2003)
Matatiella hemirosetta
pi. 52
MATATIELLALES
181
^/tRELITZIA 15 (2003)
(1 / . *
PRE/F/17250b
PRE/F/17250b
( PRE/F/ 17250b
PRE/F/17248b
*5J ipiEi^/179iib |i
M. sessilis
PRE/F/17942b
Peninsula
(Pen 41 1 Hei elo)
-a*
PRE/F/17942a Holotype
M. reducta
Aasvoelberg
(Aas411 Dic/Sph)
Holotype
PRE/F/17245a
PRE/F/20536
PRE/F/17245b
PRE/F/20536^^:
MATATIELLALES
pi. 53
Matatiella spp.
182
CV TRELITZIA 15 (2003)
GINKGOOPSIDA S.V.Meyen 1987
MATATIELLALES J.M.And. & H.M.And., ord. nov.
INCERTAE SEDIS family
SwitziantllUS J.M.And. & H.M.And., gen. nov.
Type species
Switzianthus moriformis J.M.And. & H.M.And., sp. nov.
Little Switzerland, Karoo Basin, S. Africa; Camian, Triassic.
Generic diagnosis
A ginkgoopsid male strobilus of compact conate form, with helically
arranged microsporophyll scales bearing numerous minute microsporangia
with disaccate pollen grains.
Generic characters (based on S. moriformis)
Strobilus : a simple compact cone, small to medium ( ca 25-40 mm long);
axis stout, gently curved, free end 3-6 mm; microsporophylls numer-
ous, helically arranged.
Microsporophyll. a simple scale; distal lamina broadly triangular, margin
entire, outer rim with fine radiating striae; microsporangia apparently
adaxial, numerous, closely packed.
Microsporangium: minute (0.05-0.15 mm diam.), shape uncertain.
Pollen: disaccate.
Etymology
Switzianthus— after the type locality. Little Switzerland.
Global range: 2 spp., Gondwana, Tr. (CRN).
First & last: the 2 Molteno species described here.
Gondwana Triassic occurrence
SAf— Karoo Basin, 3TCs (54 indivs).
Aus— Clarence-Moreton Basin, 1 TC (?indivs).
Molteno occurrence
Frequency (F): 4 TCs (of 100 sampled in the Molteno).
Diversity (D): 2 species.
Abundance (A): 54 individuals total, very rare to extremely rare.
Lit 111 Dic/Hei: 50 indivs in 550 man-hrs cleaving (1 per 1 man-day) very rare
Win 111 Hei elo: 1
” 20
” (i ’
2
’ )
Mat 1 1 1 Die dub: 1
” 65
” (1 ’
6
’ ) extr. rare
Note that the figures for Lit 111 refer only to curated individuals, a
selection of the best seen at the site. The rate of yield of Switzianthus at this
site may, in reality, be as high as 10 per 1 man-day.
Affiliated organs
Female strobilus: see discussion below.
Foliage: Dejerseya— Grade 3 (Cut. corn, Mut. occ.).
Classification & comparison
Suprageneric classification (fam. Incertae/Matatiellales)
The putative affiliates, Switzianthus and Dejerseya , present conflicting
evidence with regard to their classification. Switzianthus is very like
Androstrobus, which is cycadalean. while Dejerseya , in leaf shape, vena-
tion and cuticular features, is typically ginkgoopsid. However, the cone
scales of Switzianthus have a cuticular structure typical of Dejerseya and
unlike any cycads known to us. Furthermore, the affiliated female structure
(known from Dinmore, Australia, see box opposite) also points to a gink-
goopsid connection, in having similarities with Matatiella.
Considering the comparison with the ovulate organ Matatiella , and
since Dejerseya , in its pinnate form, comes close to certain species of
Kurtziana (see p. 177), supposed affiliate of Matatiella. we place
Switzianthus in the order Matatiellales. The sum of differences between the
plant-genera Dejerseya/Switzianthus and Matatiella! Kurtziana are suffi-
ciently great, however, to suggest that they should belong in distinct fami-
lies. We leave the family for Switzianthus unnamed.
Intergeneric comparison (Gondwana Triassic)
Switzianthus is close to Androstrobus in being cone-like and in bearing
numerous helically arranged microsporangiate scales. However, typical
Androstrobus scales (p. 1 36) are heeled, woody and bear abaxial microspo-
rangia with monosulcate grains, while S. moriformis scales are unheeled,
apparently fleshy, and bear microsporangia (that are seemingly adaxial)
with disaccate pollen grains.
Reconstructions
S. moriformis (Lit 111): Only a few cones in the 50-specimen S. moriformis
reference palaeodeme (Lit 111) show a few relatively clear scales, the upper
part with striations and the lower with irregularly circular depressions rep-
resenting microsporangia or at least their attachment sites [pi. 55( 1-6)]. The
evidence suggests that the microsporangia are adaxial and that most scales,
being outer views, show no details of the pollen sacs. Based on the depres-
sions, the microsporangia are very small, 0.05-0.15 mm in diameter at their
base. Their shape and length are unknown. Pollen has been isolated but no
sporangial sacs were observed. On specimen BP/2/2154 (not illustrated), a
central depression indicates the axis of the cone.
S. crispiformis (Aas 41 1 ): At Aas 41 1, the preservation is in 3D and no car-
bonaceous material is preserved. Circular depressions are visible on some
of the scales [pi. 57(5)]. The scale peduncle has not been observed in either
of the two available specimens. In the holotype, PRE/F/12935a,b, no axis
is preserved and its diameter has been estimated from the overall width of
the cone.
Switzianthus
MATATIELLALES
c/tRELITZIA 15 (2003)
183
Evidence for affiliation of organs
Mutual occurrence
Although Switzianthus is found in only two of the five TCs yielding
Dejerseya (Tab. 45), there is a strong case for affiliation at one of these,
namely Lit 1 11. In that TC, Dejerseya (at 20%) is a co-dominant member
of the diverse assemblage, and Switzianthus is the most common of the
seven male-fruit genera identified (Tab. 27). At Win 111, where a single
specimen of Switzianthus occurs, Dejerseya (at 10%) is a particularly
common element of the assemblage. At Aas 411, two specimens of
Switzianthus are present but no Dejerseya has been recognised. However,
the 15 specimens identified as Kurtziana sp. K could conceivably be a
species of a deeply pinnate form of Dejerseya (see Tab. 44).
Cuticular correspondence
Based on material from Lit 111, the cuticles of Switzianthus and the
leaf genus Dejerseya are so closely similar that it is most likely that the
two organs derive from the same natural taxon (see pp. 184, 186).
The Androstrobus puzzle
The male cone Androstrobus has been affiliated with cycad fronds by
Harris (1964). In gross morphology, Androstrobus is very similar to
Switzianthus. Thus, in those Molteno TCs where cuticle is not preserved,
it is unsure whether we are dealing with Androstrobus or Switzianthus.
We have described two specimens from Aas 411 as Switzianthus even
though the presence at that TC of cycad leaves ( Pseudoctenis , Tab. 45)
suggests the possibility that they may belong in Androstrobus.
Comparisons beyond the Molteno
Australia
Dejerseya is known from several localities in Queensland and
Tasmania. One of us (HMA) visited the Dinmore locality at Ipswich,
Queensland, in 1988 and observed numerous large slabs with bedding-
plane assemblages consisting exclusively of simple to pinnate Dejerseya
leaves and accompanying fertile material. What we regarded at that time
as the female strobilus (And. & And. 1989, p. 258) is now identified as
Switzianthus. In size and shape, the Dinmore microsporophylls (tf. 1
below) are close to Switzianthus from Lit 111. The differences in clarity
of scale outline possibly lie in the type of preservation, being carbona-
ceous compressions without well-demarcated scales at Lit 111 and
impressions with clear scales at Dinmore.
On the same bedding planes at Dinmore there are numerous four-
lobed structures (tf. 2 below) of a form not known from the Molteno.
These are probably the detached female megasporophylls of Dejerseya
and show some resemblance to Fanerotheca and Matatiella.
assemblages
(taphocoenoses)
Dejerseya
Pseudoctenis
Jeanjacquesia
Ctenis
Moltenia
Q, Switzianthus
Q. Androstrobus
1
Birlll Sph 2spp
_
i! i! -I -
-! -
2
Gre 121 Hei elo
2
3
Boe 111 Dic/Hei
-
i\ i -! "
-! -
4
Vin 111 Die odo
-
3’ -1 -i -
-1 -
5
Ela 111 Die odo
-
2> -
-1 -
6
Kra311 Die odo
-1 1
7
Wal 111 Die odo
-
2; 1: -1 -
-[ -
8
Kon 222 Die odo
-
3 j -: -
-j -
9
Kon 221 Ast 2spp
-
25 4 2 .
-
10
Kon 211
-
21 rJ -
-i -
11
Kon 111 Die odo
-
5 - -
.
12
Pen 321 Dic/Ris
-
1!
-! 1
13
Pen 221 Dic/Equ
-
2: ■>, -
14
Pen 431
-
5 - -
15
Kap 111 Dic/Ris
-
40 1 -1 -
-1 -
16
Win 111 Hei elo
10
"I 1 -1 -
1 -
17
Hla 211 Die 3spp
-
3! -! -! 2
-! -
18
" 212
-
1-1-4
-
19
" 213 Die elo
-
27; 4 -i 7
.! -
20
Umk 111 Die 2spp
-
1 -TIT 41
"I -
21
San 111 Die era
-
1 - - -
22
Mat 111 Die dub
-
12 4 -! -
l! -
23
Lit 111 Dic/Hei
20
48 6 -; 1
50 -
24
Aas 611 Hei elo
20
-} 4 -1 -
_l
25
" 211 Hei elo
14
.1 -1 .I :
- -
26
" 411 Die sph
-
19 - -
2 -
Total TCs
5
21. 3l 2, 5
4 2
Total indivs
%
%j 8 3i 55
54 2
Tab. 45. Switzianthus! Dejerseya,
Molteno occurrence
Comparisons beyond Gondwana Triassic
Laurasia Triassic— Switzianthus has some similarity to the male
cone of Bernettia phialophora from the Rhaeto-Liassic of Greenland,
which Harris (1935) attributed to Sphenobaiera spectabilis on the
basis of repeated co-occurrence and cuticular structure. Interestingly,
Harris (1935) noted that Bernettia looked very like a cycad male
cone, but reasoned that the Ginkgoales could include a wide range of
reproductive organs. Bernettia differs from Switzianthus in yielding
nondisaccate grains. Bernettia phialophora was transferred to
Androstrobus phialophora by Van Konijnenburg-Van Cittert (1993).
GONDWANA TRIASSIC, GEOSTRAT DISTRIBUTION
MATATIELLALES
Switzianthus
184
& TRELITZIA 15 (2003)
Cuticles
Potential sample: Lit 111, 50 indivs.
Macerated (this work): 3 indivs.
Presen’ation grade: Grade 5 (excellent), all features clear, large pieces.
Diagnostic characters: cells isodiametric, walls gently curved, nonpapil-
late; stomata hypostomatic, nonaligned; subsidiary cells irregular
(brachypary)actinocytic, strongly cutinised, walls thickened, florin
ring nonlappetate, guard cells probably elliptic.
Comment: both upper and lower cuticle are clearly present, but it cannot
be determined which is which.
Significance:
Classification — the cuticles, in being so similar to Dejerseya, point to
Switzianthus belonging to the Ginkgoopsida.
Affiliations— The cuticular correspondence between this male stro-
bilus and the leaf Dejerseya (pp. 186, 187; also see And. & And. 1989. p.
258) is so close that they most probably came from the same parent-plant
genus.
Pollen
In an attempt to better classify this cone, some of the scales from Lit
1 1 1 were macerated and. most unexpectedly, disaccate pollen in aggregat-
ed masses was found. The pollen occurs in definite clusters, but no sac-like
structure has been isolated. It remains uncertain whether the microsporan-
gia occur adaxially (as preferred here) or abaxially. If the latter, then these
pollen cones are particularly suggestive of the Cycadopsida and Pinopsida.
The pollen sacs so characteristic of a number of the Molteno ginkgoopsid
male genera (i.e. Antevsia, Stachyopitys, Pteruchus and Kannaskoppian-
thus) are not at all evident. However, the cuticle structure and nature of the
cutinised stoma with lappetate subsidiary cells is typical for Ginkgoopsida
and tends to confirm the affiliation with Dejerseya leaves. The cones from
the three further localities ( Aas 411. Mat 111, Win 111) do not have cuticle
for comparison.
S. moriformis
Lit 111
S. moriformis
Lit 111
?upper cuticle
?lower cuticle
Switzianthus
MATATIELLALES
d/TRELITZIA 15 (2003)
185
Switzianthus moriformis j.M.And. & H.M.And., sp. nov.
Holotype
Specimen : BP/2/2165 a,b; pi. 54(6, 8).
Assemblage (TC): Lit 111 Dic/Hei, Little Switzerland.
Preservation: complete cone with free axis, part and counterpart, longitu-
dinal external view, outline of scales unclear; compression, in thinly
laminated carbonaceous (good cuticle), dark grey shale with moderate
cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 50 indivs (all intact), pis 54, 55.
Sister palaeodemes— nil.
Specific diagnosis
A. Switzianthus species of small size with indistinctly defined, possibly
fleshy, microsporophyll scales.
Specific characters
Strobilus: cone small (ca 35 mm long); microsporophylls ca 8-10 per gyre.
Microsporophyll: scale possibly fleshy (ca 3 X 2.4 mm), margins of distal
lamina indistinct.
Etymology
moriformis— morus (Lat.), mulberry.
Comment & comparison
S. moriformis is based on a single particularly well represented
palaeodeme of 50 individuals, eight of which are illustrated adjacent (tfs
2-9). Though the compression material does not show the scale lamina in
clear definition, it has yielded excellently preserved cuticle and in situ
pollen grains. The species gives a fleshy appearance, with the scales seem-
ingly coherent or semicoherent.
Switzianthus crispiformis J.M.And. & H.M.And., sp nov.
Holotype
Specimen: PRE/F/12935 a,b; pi. 57(1-5).
Assemblage (TC): Aas411 Dic/Sph, Aasvoelberg.
Preservation: almost complete cone without free axis, longitudinal exter-
nal view; 3D impression, in thinly laminated, strongly baked, yellow-
ish grey shale with very good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 2 indivs (1 intact, 1 partial), pi. 57(1-5).
Sister palaeodemes— 2 (both listed)
Mat 111 Die dub: 1 indiv. (intact with stalk).
Win 1 1 1 Hei elo: 1 indiv. (intact without stalk).
Specific diagnosis
A Switzianthus species of moderate size with sharply defined, possibly
woody, microsporophyll scales.
Specific characters
Strobilus: cone medium-sized (up to perhaps 50 mm long); microsporo-
phylls ca 6 per gyre.
Microsporophyll: scale possibly woody (ca 5x6 mm), margins of distal
lamina sharply defined.
Etymology
crispiformis— crispus (Lat.), crisp.
Comment & comparison
Although based on a reference palaeodeme of only two specimens, S.
crispiformis is confidently recognised as a distinct species. It is larger than
S. moriformis (from Lit 111) and the scales are very much more sharply
defined. The single specimens (tfs 12, 13 adjacent) from Mat 111 and Win
1 1 1 are included here with much reservation: they may equally well repre-
sent two additional species.
It is quite possible that S. crispiformis should be transferred to the
cycadopsid genus Androstrobus: in view of this uncertainty the generic
diagnosis of Switzianthus is based exclusively on S. moriformis.
MATATIELLALES
Switzianthus
186
Dejersey a R.Herbst 1977
Type species
Dejerseya lunensis (Johnston) And. & And. 1989.
Southport, near Ida Bay, Tasmania, Australia; Camian, Triassic.
Generic concept
A ginkgoopsid leaf with linear to elliptic, simple to deeply lobed lamina.
Generic characters (Molteno Formation)
Leaf, medium-sized, linear to broadly elliptic; lamina margin ranging from
entire to gently sinuate to lobed to almost pinnate; apex obtuse; petiole
indistinct; venation pattern dependent on degree of lobing, midrib
prominent; secondary veins arching at 30-60°, forking several times.
Cuticle : see And. & And. (1989, p. 258), this vol., tfs 12-15 below.
Eponymy
Dejerseya— for Dr N.J. de Jersey, Queensland palynologist and palaeo-
botanist.
Global range: 1 sp., Gondwana Tr. (CRN).
First & last: Dejerseya lunensis (And. & And. 1989, p. 258); Molteno Fm.,
S. Africa; Blackstone Fm., Ipswich Basin, Australia.
Gondwana Triassic occurrence
Frequency (F): 7 degree squares (of the 84 across Gondwana).
Ubiquity (U): 2 continents (of 5 comprising Gondwana).
Diversity (D): 1 species.
Abundance (A): 11% (the norm in Molteno TCs).
Longevity (L): 2 myrs (Lower Camian).
Colonisation success : FUDAL rating 7/2/1/11/2 = 23.
Limited success (Grade 2): Dejerseya was the 15th most prominent
genus in the Gondwana Triassic; it was relatively abundant where it
occurred, but was infrequent and included only a single very variable
species.
Endemism: the genus, as currently known, shows a clearly disjunct distri-
bution both in Gondwana (Karoo Basin, Clarence-Moreton Basin,
Tasmania) and within the Karoo (three widely separated TCs).
Molteno occurrence
Frequency (F); 5 TCs (of 100 sampled in the Molteno).
Diversity (D): 1 species (with much intraspecific variation).
Abundance (A): co-dominant (20%) in 2 TCs; occasional to abundant
(2-10%) in 2 TCs, rare «1%) in 1 TC.
Habit: possibly a small spreading tree.
Preferred habitat: Heidiphyllum thicket.
Affiliated organs
Female strobilus: see discussion on p. 1 82.
Male strobilus: Switzianthus— Grade 3 (Cut. cor., Mut. occ.).
Classification & comparison
Intergeneric comparison
Dejerseya resembles some Dicroidium species, e.g. D. coriacium ,
in its essentially nonpinnate laminae and the venation, but differs in its
unforked frond. The cuticle, with variously distant to proximate lappets
and distinctively striate guard cells, distinguishes Dejerseya from all
other Gondwana ginkgoopsid genera.
Interspecific comparison
At present, Dejerseya is regarded as a single, highly polymorphic
species. It is known from five TCs in the Molteno Fm. and seven in
Australia.
D. lunensis
Lit 111 PRE/F/10545
/S /-rtf'’
if
w// ' '
tfs 1-15 from
And & And., 1989
Dejerseya
MATATIELLALES
TRELITZIA 15 (2003)
187
MATATIELLALES
Dejerseya
188
TRELITZIA 15 (2003)
Little Switzerland
(Lit 111 Dic/Hei)
Holotype
Switzianthus moriformis
pi. 54
MATATIELLALES
189
d^TRELITZIA 15 (2003)
MATATIELLALES
pi. 55
S witzianth us moriformis
190
& TRELITZIA 15 (2003)
PRE/F/5933
X100
Little Switzerland
(Lit 111 Dic/Hei)
BP/2/2160
X250
BP/2/2160
X100
BP/2/2160
BP/2/2160 l-
BP/2/2160
X100
PRE/F/5933
PRE/F/5933
X250
S witzianth us moriformis
pi. 56
MATATIELLALES
^/trELITZIA 15 (2003)
191
Winnaarspruit
(Win 111 Hei elo)
PRE/F/12935a
Holotype
Aasvoelberg
(Aas 411 Dic/Sph)
Matatiele
(Mat 111 Die dub)
/ PRE/F/12935a ,
MATATIELLALES
pi. 57
Switzianthus crispiformis
192
d^TRELITZIA 15 (2003)
GINKGOOPSIDA S.V.Meyen 1984
GINKGOALES Engl. 1897
AVATIACEAE J.M.And. & H.M.And., fam. nov.
Avatia J.M.And. & H.M.And.. gen. nov.
Type species
Avatia bifurcata J.M.And. & H.M.And., sp. nov.
Birds River, Karoo Basin, S. Africa; Carnian, Triassic.
Generic diagnosis
A ginkgoopsid strobilus consisting of a simple, once-forked axis bear-
ing a pair of megasporophylls each consisting of a single, erect, leafy, shal-
lowly lobed, palmate, multiovulate head.
Generic characters
Strobilus : simple, lax, bilaterally symmetrical, small ( ca 20-25 mm long);
axis gracile, elongate, distally forked; megasporophylls a single distal
pair.
Megasporophyli. simple, pedunculate; multiovulate heads palmate (6x8
mm), shallowly 4-6-lobed; ovules/seeds readily dehisced, one per
lobe.
Ovule/seed : irregularly oval to elliptical (ca 4x2 mm), narrowly winged,
with characteristic elliptical scars (possibly fungal).
Etymology
Avatia— Avis (Lat.), bird, after the type locality Birds River.
Global range: 1 sp., Gondwana, Tr. (CRN).
First & last: the Molteno species described here.
Gondwana Triassic occurrence
SAf— Karoo Basin. 6 TCs (>110 indivs).
Molteno occurrence
Frequency (F): 6 TCs (of 100 sampled in Molteno).
Diversity (D): 1 species.
Abundance (A): >110 individuals, rare to very rare.
Bir 311 Hei/Sph: 1 indiv in 2 man-hrs cleaving ( 5 per 1 man-day) rare
Nuw 1 1 1 Die zub:
5 ”
” 21
” ( 3 •' 1 ”
Aas 411 Dic/Sph:
38 ”
” 512
” (>1 ” 1 ”
Bir 1 1 1 Sph 2spp:
66 ”
” 550
” (>1 ” 1 ”
Tin 121 Sph 2spp:
1 ”
” 5
” ( 2 ” 1
Lut 311 Hei elo:
3 ”
” 50 ”
” ( 1 ” 2 ”
The figures for Bir 1 1 1 and Aas 411 refer only to curated individuals —
a selection of the best specimens from the site. The rate of yield of Avatia
at these two TCs is, in reality, much higher.
Affiliated organs
Male strobilus: Eosteria— Grade 3 (Kin. reinf., Mut. occ.).
Foliage: Ginkgoites—G rade 2 (Mut. occ.).
Classification & comparison
Suprageneric classification (Avatiaceae/Ginkgoales)
In view of the generalised similarities between Avatia and the extant
Ginkgo female strobilus (see box opposite), the probable affiliation with
Eosteria (comparable to the male strobilus of modern Ginkgo), and the
most-likely affiliation with Ginkgoites leaves, we include Avatia in the
order Ginkgoales. However, the difference between Avatia and Ginkgo
(ovulate strobilus) is sufficiently great to warrant placing the former in a
new family (Avatiaceae). It is essentially in the number of ovules per
ovuliferous head that the divergence in morphology lies.
Intergeneric comparison (Gondwana Triassic)
An intriguing and varying range of similarities occur between Avatia
and the other ovulate ginkgoopsid genera found in the Molteno.
Fanerotheca , with the winged seed Feruglioa, bears 4-lobed cupules, but
these occur in multiple pairs along megasporophylls attached to a full stro-
bilus. Umkomasia , in some species, bears once-forked megasporophylls,
but these do not have palmate heads. Peltaspennum is characterised by
peltate, multilobed ovulate heads that readily detach. Matatiella has a sim-
ilar palmate megasporophyli, but it is deeply lobed and attached helically
to an axis. Hamshawvia has a once-forked strobilus with a pair of megas-
porophylls, but these are fleshy and with 8 to 16 small ovules.
Reconstructions
We have attempted no more than R2-3 reconstructions of Avatia from
the reference palaeodeme, Bir 111, or the sister palaeodemes, Aas 411 and
Lut 311 (see above). While there is good evidence that the seeds shown are
affiliated with Avatia, it is unknown whether they are attached adaxially,
abaxially, or enclosed within the megasporophyli lobe.
Avatia
GINKGOALES
d^TRELITZIA 15 (2003)
193
Evidence for affiliation of organs
Kindred reinforcement
Avatia resembles the extant Ginkgo strobilus in its typical pair of
ovules (tfs 1-16 opposite). There are also similarities with some of the
anomalous forms of Ginkgo strobili (tfs 18, 19 opposite) as reported by
Seward & Gowan (1900, fig. 37) and Krassilov (1972, fig. la, b). See Hara
(1997) for a recent discussion and further examples of anomalous strobili.
However, the fleshy fruit and hard pip of Ginkgo biloba is quite unlike the
palmate head and winged seed of Avatia.
Mutual occurrence
Foliage— Ginkgoites occurs frequently at Bir 111 and Aas 41 1 and is
the most likely nominee for affiliation with Avatia (Grade 2). It is absent
from Lut 3 1 1 and Nuw 111 (Tab. 46). If Sphenobaiera was not already pre-
occupied (Grade 3 affiliation with Hamshawvia ), it would be a more or less
equal candidate for affiliation with Avatia. The immature female strobilus
attached to the short shoot with Sphenobaiera [p. 223, tf. 10; pi. 7 1 ( 1—4)],
while difficult to identify, could also be Avatia. No other gymnospermous
foliage in the Molteno shows a pattern of occurrence suggesting any likely
link with Avatia.
Male—Eosteria ; see discussion under that genus (p. 196).
Seeds— A distinctive narrowly winged seed (pi. 59) co-occurs with
Avatia in the three TCs listed below, suggesting an affiliation of Grade 3
reliability. This affiliation is supported by the presence of the elliptical
scars, possibly of fungal origin, that occur on both the seeds and the megas-
porophyll lobes.
Bir 111— Numerous seeds occur either scattered or in dense clusters, pi.
58(7), and in several cases, pi. 58(3, 4), they lie in close proximity to
Avatia megasporophylls, but, unfortunately, are never found attached.
The elliptical scars which are very evident on the seeds, pi. 59(4-9),
are seen also in one of the illustrated palmate heads, pi. 59(2).
Aas 411 — One slab, PRE/F/20682, pi. 58(10), has two Avatia individuals
with the affiliated seeds in close proximity as if they may have been
attached in life.
Nuw 777— Three of the five megasporophyll-bearing slabs show several
dispersed seeds. None of these are attached or bear the characteristic
elliptical scars. Numerous slabs show only dispersed seeds.
assemblages
(taphocoenosis)
Tel 111
Vin 111
Ela 111
Lut 311 Hei elo
Tin 121 Sph 2spp
Wal 111 Die odo
Kon 223 j;
222
Nuw 111 Die zub
211 Die 2spp
Maz 111 Die era
211 Hei/Dic
San 111 Die era
Mng 111 Pic 2spp
Mat 111 Die dub
Lit 111 Dic/Hei
Aas 311 Hei elo
411 Dic/Sph
Total TCs
Total indivs
Tab. 46. Avatia/Ginkgoites, Molteno occurrence
Abundance: individuals of Avatia are abundant at both Bir 111 and Aas 411.
In these TCs only the number of curated specimens is recorded. Many further
isolated individuals occur on other slabs in the collection.
Comparisons beyond Gondwana Triassic
The ovulate organs from outside the Gondwana Triassic most sim-
ilar to Avatia are the glossopterid genera Rigbya and Lidgettonia from
the Gondwana Permian (And. & And. 1985, pp. 127-136). These both
have lobed megasporophyll heads, apparently bearing winged seeds.
Though neither are forked like Avatia, either could have evolved over a
geological period to look similar. The seeds affiliated here with Avatia
are, in hand specimens, closely similar to those that have been affiliat-
ed with another putative glossopterid ovulate fruit, Arberia (And. &
And. 1985, pp. 128-131).
Zhou (1991, 1997) reviewed ginkgoalean female megafossils
attached or affiliated with Ginkgo fossil leaves. None of these, e.g.
Karkenia (Lower Cretaceous, Tico Flora, Santa Cruz Province,
Argentina), Yimaia (Middle Jurassic, Yima Fm., Henan, China) and
Umaltolepsis (Upper Jurassic & Lower Cretaceous, Bureja River Basin,
Siberia), show a resemblance to Avatia.
Meeusella proteidada
The paired heads of Meeusella were
regarded by Krassilov and Bugdaeva
as microsporangiate. However, they
could be ovulate organs and they do
show some resemblance to Avatia
from Krassilov & Bugdaeva (1988)
Lake Baikal, USSR, L. Cretaceous
Avatia bifurcata And. & And., sp. nov.
Holotype
Specimen: BP/2/5230a,b; pi. 58(1, 8).
Assemblage (TC); Bir 111 Sph 2spp, Birds River.
Preservation: virtually complete strobilus, part and counterpart; impres-
sion in thinly laminated, yellowish grey shale with very good cleavage.
Reference palaeodeme (Molteno Fm.)
Assemblage (TC): as for holotype.
Specimens: 66 indivs (most intact), pis 58(1-9), 59(1-9).
Sister palaeodemes (Molteno Fm.)— 5 (best 1 listed)
Aas 411 Dic/Sph: 38 indivs.
Specific diagnosis: as for genus.
Diagnostic characters: as for genus.
Etymology
bifurcata— with reference to the bifurcating form of the strobilus.
Classification & comparison: see under genus.
1-10 degre
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Cities j
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5
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7 ••••**;
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ano Fm.
GINKGOALES
Avatia
194
c/treliTZIA 15 (2003)
Eosteria J.M.And. & H.M.And., gen. nov.
Type species
Eosteria eosteranthus J.M.And. & H.M.And., sp. nov.
Aasvoelberg (Aas 311), Karoo Basin. S. Africa; Camian, Triassic.
Generic diagnosis
A ginkgoopsid male strobilus of linear, spicate form, with simple
microsporophylls comprising single, pedunculate microsporangiate
cupules.
Generic characters
Strobilus: simple, linear spicate. relatively lax, radially symmetrical, small
(up to ca 40 mm long); axis gracile, gently sinuous to erect;
microsporophylls numerous, helically arranged.
Microsporophyll: simple, pedunculate; peduncle gracile (ca 1 mm long);
microsporangia borne singly.
Microsporangium: cupulate, bilaterally symmetrical; closed sac ovoid (ca
2-3 mm long), opened sac relatively deeply 6-lobed.
Pollen: monosulcate, spherical with large circular germinal furrow.
Etymology
Eosteria— after a Germanic goddess Eostre ; with reference to the type
locality Aasvoelberg (Aas 311), informally called ‘Easter Egg Shale’.
Global range: 2 spp., Gondwana, Tr. (CRN).
First & last: the 2 Molteno species described here.
Gondwana Triassic occurrence
SAf— Karoo Basin, 4 TCs (27 indivs).
Molteno occurrence
Frequency (F): 4 TCs (of the 100 Molteno TCs sampled).
Diversity (D): 2 species.
Abundance (A) 27 indivs total, very rare to extremely rare.
Aas 311 Hei elo:
12 indivs in 140 man
hrs cleaving ( 1 per 1 man-day) very rare
Tel 1 1 1 Hei elo:
8 '
” 90
” (1 "1 ” )
Lut 311 Hei elo:
2 ’
” 50
” (1 ”2 ” )
Bir 1 1 1 Sph 2spp:
5 '
” 550
” ( 1 ” 1 1 ” ) extr. rare
Eosteria is an infrequent and very rare element in the Molteno.
Affiliated organs
Female strobilus: Avatia— Grade 3 (Kin. reinf., Mut. occ.).
Foliage: Ginkgoites— Grade 3 (Kin. reinf., Mut. occ.).
Classification & comparison
Suprageneric classification (Avatiaceae/Ginkgoales)
As discussed earlier (p. 192), the plant-genus comprising the affiliated
grouping of Avatia (female), Ginkgoites (foliage) and Eosteria (male), is
readily placed in the order Ginkgoales. Of all the Molteno microsporan-
giate genera, Eosteria is clearly most like the male strobilus of the extant
Ginkgo biloba (box on p. 195). In our view, both Avatia and Eosteria are
sufficiently different from their living female and male ginkgoalean coun-
terparts, however, to warrant including them in a distinct family
(Avatiaceae).
Intergeneric comparisons (Gondwana Triassic)
The other ginkgoopsid microsporangiate genera from the Molteno—
Antevsia, Switzianthus, Stachyopitys , Pteruchus and Kannaskoppianthus—
are all fundamentally different from Eosteria. None consists of a spicate
strobilus or bears cupulate microsporangia remotely like it.
Reconstructions
Strobilus
The R4 strobilus reconstructions of the two described species, E.
eosteranthus and E. telemanthus , are based on the full set of specimens in
the reference palaeodeme of each. For both species, the holotype is used as
the basis for overall size and shape, but other individuals are referred to for
details of the microsporophylls and their attachment. The full length of the
strobilus is uncertain for both species. We have chosen to show the strobili
with fully open sacs proximally and with closed sacs distally.
Microsporangia
Interpretation of the structure of the individual microsporangial sacs is
particularly difficult. In E. telemanthus , for instance, there is only one speci-
men [PRE/F/17330a‘y\ pi. 65(1-3); this lies immediately adjacent to the
holotype on the same slab) with a single opened sac that shows most of the
lobes and their margins intact. The next most fully preserved open sac is
that towards the proximal end of the holotype [PRE/F/17330a,b‘x\ pi.
64(1, 5)] and this gives a quite contradictory impression of shape. The outer
margins of the lobes angle upwards out of the sediment and are tom.
Similarly for E. eosteranthus from Aas 311: only in PRE/F/22074 [pi.
61(4)] is there an open sac with the distal lobes untorn, while other speci-
mens, e.g. PRE/F/22067, pi. 61(5-7); PRE/F/22070, pi. 60(1-5), show
open sacs with torn margins that suggest different shapes. The closed or
partially open sacs are most evident in Lut 311 (pi. 62) and Bir 1 1 1 (pi. 63).
Eosteria
GINKGOALES
TRELITZIA 15 (2003)
195
Extant Ginkgo biloba ( male strobilus)
Ginkgo biloba bears a simple linear male strobilus with numerous
spirally arranged microsporophylls on slender pedicels. The microspo-
rangia, however, are rather distinct: they consist of a closed pair of lin-
ear-oblong sacs suspended from a flattened receptacle at the end of the
pedicel; the sacs dehisce along an inner suture to release the pollen. The
dried, opened pair of sacs distort irregularly to present a variety of
shapes. Pollen is found adhering in abundance to the hirsute inner face
of the sacs.
Eosteria & G. biloba
Eosteria is closely similar to the extant Ginkgo male in its simple
strobilus with spirally arranged microsporophylls borne on slender
pedicels, but differs in its single microsporangium (not paired) and the
sac dehiscing into ca six lobes (not a single suture). Both genera yield
monocolpate pollen.
Ginkgoales
Evidence of the male strobilus of Ginkgo in the fossil record is
particularly sparse. Only four references are known to us:
• Harris & Millington (1974); Yorkshire Jurassic; 1 individual.
• Schweitzer (1977); Alborz Mountains, Rhaetic, Iran, ca 8 individuals;
Irania (Iraniales). I. hermaphroditica is an axis bearing male and
female strobili (the latter on the upper part of the axis). The
microsporophyll strobilus is very similar to the extant Ginkgo biloba
male strobilus with two pollen sacs per microsporangium. So far no
pollen grains have been found. The combination of male and female
in one structure makes Irania a unique organ and its classification an
enigma.
• Drinnan & Chambers ( 1985, 1986), Douglas (1969); Koonwarra Lake
deposit in Whitelaw road cutting, Victoria, mid-Cretaceous; four speci-
mens, two being multiple and attached to an axis.
• Rothwell & Holt ( 1997); central Alberta, Upper Cretaceous, one indi-
vidual.
pollen 4
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
GINKGOALES
Eosteria
196
Cff TRELITZIA 15 (2003)
Pollen
Three specimens, one each from Aas 311 [PRE/F/22070, pi. 60(1-5)],
Tel 111 [PRE/F/17325, pi. 65(5-7)] and Bir 111 [PRE/F/10185, pi.
63(7-9)], show in situ clusters of pollen in opened or naturally sectioned
microsporangial sacs. The most clearly preserved of these clusters is that
from Bir 1 1 1 . where, at °°80 magnification, it is quite evident that the grains
are spherical with a large rounded germinal furrow. They compare very
closely to those of extant monocolpate pollen of Ginkgo biloba (Sahashi
1997).
Evidence for affiliation of organs
A Grade 3 affiliation of Eosteria with the female strobilus Avatia and
with the foliage Ginkgoites is based on the following evidence:
Mutual occurrence (Tab. 46)
Eosteria occurs together with Ginkgoites (common, two species) and
Avatia (abundant) at Bir 1 1 1 , with only Ginkgoites (rare, one species) at Tel
111, with only Avatia (rare) at Lut 311. and with neither genus at Aas 311.
Kindred reinforcement : The extant Ginkgo biloba male strobilus (see
boxes and relevant text) is similar to Eosteria in both gross morphology
and pollen grains.
Adaptive radiation (Molteno diveristy)
It is likely that Eosteria was more diverse than the available material
(in quantity, intactness and clarity) enables us to recognise. While four
palaeodemes are represented and only two species have been defined, we
consider it more likely, in life, that four species may have been present.
The chief diagnostic characters are the shape and robustness of the
strobili, and the shape and orientation of the lobes in the fully opened
microsporangiate sacs.
The two species, each based on reasonably sized reference palaeo-
demes and deriving from the same habitat but different stratigraphic levels,
are:
E. eosteranthus— Aas 311 Hei elo (Aasvoelberg); 12 indivs
Heidiphyllum thicket; Cycle 1 (Bamboesberg Member)
E. telemanthus— Tel 111 Hei elo (Telemachus); 8 indivs
Heidiphyllum thicket; Cycle 3 (Mayaputi Member)
Eosteria telemanthus j.M.And. & H.M.And., sp. nov.
Holotype
Specimen : PRE/F/17330a,b‘x’; pi. 64(1, 2, 5).
Assemblage (TC): Tel 111 Hei elo, Telemachus Spruit.
Preservation : almost complete strobilus, part and counterpart; impression
in thickly laminated, light olive-grey shale with poor cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens : 8 indivs (1 intact, 3 partial, 4 isolated), pis 64(1-8), 65(1-7).
Sister palaeodemes— nil.
Specific diagnosis
An Eosteria species with a relatively stout strobilus and microspor-
angial sacs dehiscing radially into 6 fairly equal lobes.
Specific characteristics
Strobilus: relatively stout (length unknown).
Microsporangium: cupulate sacs dehiscing into 6 fairly equal lobes; outer
pair of proximal lobes without distinctive ribs.
Etymology
telemanthus— a male cone from Telemachus Spruit (Tel 111).
Comment & comparison
The E. telemanthus palaeodeme includes a number of very clearly pre-
served specimens. They would show up particularly well if photographed
in colour— with the rust-coloured points of pedicel attachment, for
instance, standing out sharply. The species appears distinct, as is its puta-
tive foliage affiliate, Ginkgoites telemachus , which is also well represented
(23 individuals) and exclusive to Tel 111.
Eosteria
GINKGOALES
(Ef' TRELITZIA 15 (2003)
197
Eosteria eosteranthus j.M.And. & H.M.And., sp. nov.
Holotype
Specimen : PRE/F/1 9388a, b; pi. 60(6-8).
Assemblage (TC): Aas311 Hei elo, Aasvoelberg.
Preservation : nearly complete strobilus, part and counterpart; impression,
in thickly laminated, light grey shale with moderate cleavage.
Reference palaeodeme
Assemblage ( TC): as for holotype.
Specimens: 12 indivs (3 intact, 4 partial, 5 isolated), pis 60(1-8), 61(1-7).
Sister palaeodemes— 2 (both listed)
Bir 1 1 1 Sph 2spp: 5 indivs, distinctive pollen grains; pi. 63(1-9).
Lut 311 Hei elo; 2 indivs, clear axis and pedicels; pi. 62(1-7).
Specific diagnosis
An Eosteria species with a linear, gracile strobilus and microsporan-
gial sacs dehiscing into two strongly unequal groups of three lobes.
Specific characteristics
Strobilus: linear, gracile (to ca 40 mm).
Microsporangium: cupulate sacs dehiscing into two unequal groups of 3
lobes, with a wide angle between the upper and lower sets of lobes,
outer pair of proximal lobes with distinctive rib.
Etymology
eosteranthus— a male cone from the Easter Egg Shale (i.e. Aas 311).
Comment & comparison
E. eosteranthus is distinguished from E. telemanthus in the more
gracile form of the strobilus and the clearer separation into distal and prox-
imal lobes in the opened microsporangium. The specimens from Bir 111
and Lut 311 are included in this species, but with better preserved material
they may well prove to be distinct.
"W'X,
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m situ
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PRE/F/10185
pi, 63(7-9)
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Sister palaeodeme
. . 2
w
PRE/F/22072
pi. 61(1,2)
tnis-Jt
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s
PRE/F/19388a,b
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PRE/F/22070
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PRE/F/22074
pi. 61(4)
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Reference palaeodeme
VlfrW
PRE/F/22074 /
pi. 61(4)
GINKGO ALES
Eosteria
198
d^TRELITZIA 15 (2003)
Ginkgoites Seward 1919
Type species
Ginkgoites obovata Seward 1919.
Bjuv, Scania, Sweden; Rhaetic, Triassic.
Generic concept
A ginkgoopsid leaf with distinct long petiole, deeply dichotomously
dividing fan-shaped lamina, and radiating, repeatedly forking subparallel
venation.
Generic characters (Molteno Fm.)
Attachment: unknown.
Leaf, fan-shaped with margins diverging at >90°, lamina deeply dichoto-
mously divided into several segments, with each segment variously
lobed; petiole distinct, long, gracile; veins radiating from base, repeat-
edly forking, close to well spaced, running subparallel to terminate at
distal margin.
Cuticle: see And. & And. (1989, p. 220); this vol., tfs 1^1 below.
Etymology
Ginkgoites— referring to similarity to Ginkgo.
Global range: numerous spp., Pangaea, ?Tr.-Tertiary.
Gondwana Triassic occurrence
Frequency (F): 21 degree squares (of the 84 across Gondwana).
Ubiquity (U): 4 continents (of 5 comprising Gondwana).
Diversity (D): 9 foliage species.
Abundance (A); <1% (the norm in Molteno TCs).
Longevity (L): 17 myrs (Spathian to early Camian).
Colonisation success: FUDAL rating 21/4/9/-/17 = 51.
Intermediate success (Grade 3): Ginkgoites was the 7th most promi-
nent genus in the Gondwana Triassic; it was relatively frequent,
ubiquitous, diverse and long-lived, but markedly lacking in abun-
dance.
Endemism: of the 9 species, 6 are single-assemblage endemics, 1 is a
basin endemic, 1 a continent endemic, while the last is particularly
widespread and variable and occurs throughout the realm.
Classification & comparison
The generic names Ginkgo and Ginkgoites are both in common use for
fossils (see Stewart & Rothwell 1993, p. 390). In And. & And. (1989, p.
218), we followed Harris & Millington (1974) and used Ginkgo. However,
in the absence of fruit allied to the extant Ginkgo , it seems more appropri-
ate to name the Molteno fossils Ginkgoites.
Intergeneric comparison
Gondwana Triassic— In the dichotomising nature of both lamina and
venation, Ginkgoites and Sphenobaiera are clearly the most alike of the
ginkgoopsid leaf genera. In epidermal structure, however, Ginkgoites is
also similar to Sphenobaiera , Lepidopteris and Dejerseya, but can be dis-
tinquished by the finer details of the cuticles. The cuticles of all three
species of Ginkgoites occurring at Lit 111 (Tab. 46) are illustrated and
described in And. & And. ( 1 989).
The leaf species that in gross morphology is closest to the extant
Ginkgo biloba has here been transferred to the new genus Paraginkgo (p.
208). The cuticular details of this species, P. antarctica, are markedly dis-
tinct from both the extant Ginkgo biloba and fossil Ginkgoites cuticles.
Interspecific comparison
The nine species of Ginkgoites recognised from the Gondwana
Triassic fall readily within the compass of the genus. The cuticle, howev-
er, is known for only three of these species, so confirmation is not at hand
from this valuable source. The six Molteno Ginkgoites species are based on
distinct, nonoverlapping reference palaeodemes. While most species, as
known, are single-assemblage endemics, one species (G. matatiensis ) is
highly polymorphic and occurs in 16 Molteno TCs.
Molteno occurrence
Frequency (F): 19 TCs (of 100 sampled in the Molteno).
Diversity (D): 6 species.
Abundance (A): occasional to common (1-5%) in 5 TCs; rare to very
rare (<1%) in 14 TCs.
Habit: probably a tall deciduous tree.
Preferred habitat: a scattered element in riparian forest and woodland.
Affiliated organs
Female strobilus: Avatia— Grade 2 (Kin. reinf., Mut. occ.).
Male strobilus: Eosteria—G rade 3 (Kin. reinf., Mut. occ.).
Ginkgoites matatiensis
Lit 111 PRE/F/5485
Florin ring
x500
tfs 1-4 from
And. & And , 1989
Ginkgoites
GINKGOALES
^/trelitzia 15 (2003)
199
tfs 1-6 from
And. & And., 1989
Ginkgoites
generic panorama showing
all 6 Molteno species
(see fab, 46, p. 193)
G wahleckensis
G mutatiensis
r,
Wal 111
Rr PRE/F/2147b
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GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
SAm SAf Ind Ant Aus
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o: productive degree
squares
Gondwana Triassic
GINKGOALES
Ginkgoites
200
RELITZIA 15 (2003)
BP/2/5230b
Holotype
PRE/F/10736
Birds River
(Bir 111 Sph 2spp)
PRE/F/13104 PRE/F/13107b
PRE/F/10065
Aasvoelberg
(Aas411 Dic/Sph)
Lutherskop
(Lut 311 Hei elo)
BP/2/5260
Avatia bifurcata
pi. 58
GINKGOALES
TRELITZIA 15 (2003)
201
Birds River
(Bir 111 Sph 2spp)
PRE/F/10072 _/'
tzfmmm %
PRE/F/10734 *
BP/2/5393
BP/2/5397
PRE/F/10070
GINKGOALES
pi. 59
Avatia bifurcata
202
df TRELITZIA 15 (2003)
Eosteria eosteranthus
pi. 60
GINKGOALES
203
TRELITZIA 15 (2003)
PRE/F/22072
PRE/F/22069
PRE/F/22072 ^
Aasvoelberg
(Aas 311 Hei elo)
PRE/F/22074
:j>RE/F/22.067_
PRE/F/22067
PRE/F/22067
GINKGOALES
pi. 61
Eosteria eosteranthus
204
< 'if TRELITZIA 15 (2003)
«%_xl0
PRE/F/11521
Lutherskop
(Lut 3 1 1 Hei elo)
PRE/F/l 1515a X10
r-J: ;
PRE/F/l 1515a
' \
f :
PRE/F/l 1515a
i >■. xl° '
Eosteria eosteranthus
pi. 62
GINKGOALES
TRELITZIA 15 (2003)
PRE/F/15590
PRE/F/19931
PRE/F/19931
PRE/F/19931
PRE/F/19931 X5
Birds River
(Bir 111 Sph 2spp)
.y
jb
PRE/F/10051a
X40
GINKGOALES
pi. 63
Eosteria eosteranthus
c/trELITZIA 15 (2003)
• PRE/F/17324
•v *
Telemachus Spruit
(Tel 1 1 1 Hei elo)
PRE/F/17330b
X20
Holotype
Eosteria telemanthus
pi. 64
GINKGO ALES
(§P TRELITZIA 15 (2003)
207
GINKGOALES
pi. 65
Eosteria telemanthus
208
•'■'/tri; i rrziA 15 (2003)
GINKGOOPSIDA S.V.Meyen 1984
GINKGO ALES Engl. 1897
INCERTAE SEDIS family
Paraginkgo J.M.And. & H.M.And., gen. nov.
Type species
Salisburia antarctica Saporta 1882.
Sydney Basin, Australia; Ladinian, Triassic.
Generic diagnosis
A ginkgoopsid leaf with short gracile petiole, entire fan-shaped lamina
and radiating, forking, closely spaced, subparallel venation and distinct
cuticle.
Generic characters (Molteno Fm.)
Attachment: unknown.
Leaf, small ( ca 30 X 60 mm), broadly fan-shaped with margins diverging
at >90°, lamina entire to sinuate with occasional shallow clefts; petiole
distinct, short, gracile; veins radiating from base, repeatedly forking,
closely spaced and subparallel (20 per 10 mm at distal margin).
Cuticle : see And. & And. (1989, p. 220); this vol., tfs 1—4 opposite.
Etymology
Paraginkgo— with reference to the similarity in gross form to extant
Ginkgo biloba leaves.
Global Range: 1 sp., Gondwana, U.Tr. (LAD-CRN).
First: Paraginkgo antarctica (And. & And. 1989, p. 540); Culvida Sdst.,
Mt Ernest (C62), Canning Basin, Western Australia.
Last: the Molteno species described here.
Gondwana Triassic occurrence (after And. & And. 1989)
Frequency (F): 5 degree squares (of the 84 across Gondwana).
Ubiquity (U): 2 continents (of 5 comprising Gondwana).
Diversity (D): 1 foliage species.
Abundance (A): <1% (the norm in Molteno TCs).
Longevity (L): 14 myrs (Spathian-Camian).
Colonisation success: FUDAL rating 5/2/1/—/ 1 4 = 22.
Limited success (Grade 2): Paraginkgo was the 16th most prominent
genus in the Gondwana Triassic; it was widespread (though disjunct)
and long-lived, but lacking in frequency, abundance and diversity.
Endemism: the single species had a disjunct distribution (Karoo Basin
and Australia).
Molteno occurrence
Frequency (F): 2 TCs (of 100 sampled in the Molteno).
Diversity (D): 1 species.
Abundance (A): very rare (<1%) in both TCs.
Habit: probably a shrub.
Preferred habitat: Dicroidium riparian forest.
Affiliated organs: unknown.
Classification & comparison
The systematic position of P. antarctica is problematic. In gross form
it closely resembles the extant Ginkgo biloba. However, the cuticle of this
extant species does not appear to be closely related to either the fossil
Ginkgoites species or to P. antarctica. The cuticle of P. antarctica is dis-
tinctly different from that of G. matatiensis and G. telemachus from Lit 1 1 1
which have lappetate subsidiary cells and no striae on the guard cells. Nor
does it resemble the cuticle of the bulk of other fossil Ginkgoites species
from around the globe. These latter form an apparently natural group to
which G. matatiensis and G. teleinachus belong.
Paraginkgo antarctica (Saporta) J.M.And. & H.M.And.,
comb nov.
Holotype
Specimen: catalogue number and repository not in literature. See Saporta
[1882, tf. 1(1)] and And. & And. [1989, pi. 320(1)].
Assemblage/locality: Sydney region, Wianamatta Grp., Sydney Basin,
Ladinian, Triassic.
Reference assemblage & palaeodeme
Assemblage: Lit 111 Dic/Hei, Little Switzerland.
Specimens: 40 individuals (see And. & And. 1989, pi. 113-118).
Cuticle: well preserved.
Sister palaeodeme
San 1 1 1 Die era: 3 indivs.
Specific diagnosis: as for genus.
Specific characters: as for genus.
Etymology
antarctica— probably for the southern occurrence of the species.
Comment & comparison: as for genus.
Paraginkgo
GINKGOALES
c/tRELITZIA 15 (2003)
209
Cuticles
Potential sample : Lit 111, 40 indivs.
Macerated (this work): Lit 111, 3 indivs.
Preservation grade : Grade 5.
Diagnostic characters: with nonlappetate subsidiary cells and guard
cells with strong radial striae.
Comment: Paraginkgo has been separated from Ginkgoites primarily on
the basis of its very distinctive cuticle.
Significance:
Classification— of no aid beyond suggesting the class Ginkgoopsida.
Affiliations— no foliage species with known cuticle matches this.
P. antarctica
3
tfs l^t
from And & And., 1989
2
lower
BP/2/1859
xlOO
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
GINKGOALES
Paraginkgo
210
TREL1TZIA 15 (2003)
GINKGOOPSIDA S.V.Meyen 1984
HAMSHAWVIALES J.M.And. & H.M.And., ord. nov.
HAMSHAWVIACEAE J.M.And. & H.M.And., fam. nov.
Hamshawvia J.M.And. & H.M.And., gen. nov.
Type species
Hamshawvia baccata J.M.And. & H.M.And., sp. nov.
Umkomaas Valley. Karoo Basin, S. Africa; Camian, Triassic.
Generic diagnosis
A ginkgoopsid strobilus consisting of a simple, once-forked axis, bear-
ing a pair of megasporophylls each consisting of single, erect, fleshy,
rounded multiovulate lamina.
Generic characters (based on the 4 Molteno species)
Attachment : Strobili borne on a short shoot with several Sphenobaiera
leaves.
Strobilus'. simple, lax, bilaterally symmetrical, small (ca 1 5—40 mm long);
axis relatively gracile, short to elongate, distally forked; megasporo-
phylls a single distal pair.
Megasporophyl! : simple, pedunculate; ovuliferous lamina rounded to
cordiform, apparently fleshy, bilaterally symmetrical, flattened dor-
siventrally, sometimes apparently longitudinally folded; ovules/seeds
adaxial, embedded, ca 8-20 in number, arranged radially to semiradi-
ally or bilaterally on either side of a median vein or receptacle.
Ovule/seed: small (up to ca 1 mm long), generally elliptical; with a slight-
ly elongate central marking (?embryo).
Eponymy
Hamshawvia— in honour of Hugh Hamshaw Thomas, one of the pioneer
collectors from the famous Umkomaas Valley (Umk 111) locality.
Global range: 5 spp., GondwanaTr. (ANS-CRN).
First: Hamshawvia sp. (Holmes 1995); Basin Creek Fm., Nymboida,
N.S.W., Australia.
Last: the 4 Molteno species described here.
Gondwana Triassic occurrence
S Am— Argentina, 1 TC (1 indiv.).
SAf— Karoo Basin, 4 TCs (24 indivs).
Aus— Nymboida Sub-basin, 1 TC (2 indivs).
Molteno occurrence
Frequency (F): 4 TCs (of 100 sampled in the Molteno).
Diversity (D); 4 species.
Abundance (A): 24 indivs total, very rare to extremely rare.
Aas 411 Dic/Sph: 14 indivs in 512 man-hrs (1 per 4 man-days) very rare
Lut 311 Hei elo:
1 ”
” 50
(1
5
)
Umk 1 1 1 Die 2spp:
7 ”
” 400
(1
6
) extremely rare
Lit 111 Dic/Hei:
2 ”
” 550
(1
27
)
Hamshawvia is an
nfrequent and very
rare element in the Molteno.
Affiliated organs (for details see p. 213)
Male strobilus: Stachyopitys— Grade 4 (Mut. occ., Kin. reinf.).
Foliage: Sphenobaiera— Grade 4/5 (Mut. occ., Kin. reinf., Org. att., Cut.
corr.).
Classification & comparison
Suprageneric classification (Hamshawviaceae/Hamshawviales)
Without considering the supposed affiliated organs of Hamshawvia , it
would be particularly difficult to classify this genus. The simple forked
strobilus and the cuticle suggest the Ginkgoopsida, while the multiovulate
heads (megasporophylls) hint more towards the Bennettitopsida. However,
with the Sphenobaiera and Stachyopitys affiliation taken into consideration,
placement within the Ginkgoopsida— close to Dicroidium (Umkomasi-
aceae) and Ginkgoites (Avatiaceae) — is favoured. The differences between
Hamshawvia and any of the other ovulate Gondwana-Triassic ginkgoopsid
genera are profound enough to erect a new order (Hamshawviales) and
family (Hamshawviaceae) for this plant-genus.
Intergeneric comparison (Gondwana Triassic)
Of the ginkgoopsid ovulate genera in the southern Triassic kingdom,
Avatia is perhaps the nearest to Hamshawvia. Both genera consist of sim-
ple once-forked strobili attached, most likely, in the same manner as in the
extant Ginkgo biloba , to short shoots. Avatia differs in its leafy 4- or 5-
lobed heads and its readily dehisced winged seeds.
Reconstructions
Hamshawvia longipedunculata (Aas 411)
The R4 reconstruction is based on three specimens in particular:
PRE/F/12932b (p. 215, tf. 2) for the forked pair of megasporophylls;
PRE/F/21778 (p. 215, tf. 1) for the arrangement of seeds in the apparently
fleshy megasporophyll head; and PRE/F/21779 (p. 215. tf. 4) for the elon-
gate peduncle.
A characteristic feature seen in a few specimens, most clearly in
PRE/F/21438 (p. 215, tf. 5), is the radiating pattern developed around each
seed. This feature is interpreted as the result of postmaturity shrinkage. We
suggest that in life the megasporophyll was a fleshy structure with embed-
ded seeds, well adapted for dispersal by fructivores.
Hamshawvia baccata (Umk 111)
The R4 reconstruction is a combination of two specimens:
PRE/F/1 124 (p. 214, tf.l) for the overall form of the strobilus and for the
arrangement of seeds embedded within the inner face of the megasporo-
phyll heads; and BP/2/9877b (p. 214, tf. 3) for the folding of the megas-
porophyll about a central rib.
In tfs 4a-c (above) our interpretation of the maturing megasporophylls
as they unfold is shown. The mature heads were apparently flattened berry-
like structures, but in the absence of shrinkage patterns, are seen as less
fleshy than in H. longipedunculata.
Fertile short shoot (Aas 411)
The Hamshawvia! Sphenobaiera reconstruction (tf. 1 opposite) is
based on PRE/F/12823a,b [pi. 7 1 ( 1-4)], which bears a single (?)immature
individual of Hamshawvia together with four Sphenobaiera schenckii
leaves terminally on a (?)short shoot. In the reconstruction, we show three
mature H. longipedunculata strobili within a cluster of several 5. schenckii
leaves as might have been expected in life.
Hamshawvia
HAMSHAWVIALES
d^TRELITZIA 15 (2003)
211
Hamshawvia longipedunculata \ x
ovulate organs attached to a stem
bearing Sphenobaiera schenckii leaves
(see p. 223)
Ovule/seeds
On the specimens available, the ovules/seeds are rarely seen or are
barely evident. We assume that they were well embedded within the
(?)fleshy fruit. The seeds are interpreted further as occurring in a single
plane rather than radiating hemispherically from the median axis since
some specimens from Umk 111, e.g. BP/2/9877b and BP/2/1125, show the
folded nature of the head about the axis. However, this is not seen in the
Aas 411 specimens which in the strongly wrinkled pattern indicate that the
fruits were very fleshy and possibly more spherical. In PRE/F/21438a [pi.
70(7)], the ovules are particularly clear owing to the differential shrinkage
of the surrounding tissue.
So far no ovules/seeds have been isolated by maceration. Specimen
BP/2/1125, pi. 67(3-5), did yield ovoid opaque structures, but they are
more likely to be resin bodies. Compare these with the more rounded
(?)resin bodies from BP/2/1124, pi. 67(10, 11).
Cuticles
Potential sample'. Lit 111, 2 indivs.; Umk 111,7 indivs.
Macerated (this work): Lit 111, 2 indivs; Umk 111,2 indivs.
Preservation grade : Lit 111, Grade 4, features clear, good pieces; Umk
111, Grade 3, features visible, fair pieces.
Diagnostic characters : cells isodiametric to linear oblong, walls gently
curved; papillae single, central, medium bold, restricted to one surface;
hypostomatic, stoma nonaligned; subsidiary cells brachypara(actino)-
cytic, moderately cutinised, radial walls thickened; guard cells elliptic;
florin ring lappetate.
Comment', possible resin bodies noted adjacent.
Significance:
Classification— The cuticular features support the placement of
Hamshawvia in the Ginkgoopsida which often have isodiametric, papillate
cells, and lappetate subsidiary cells.
Affiliations— The Hamshawvia cuticle from Lit 111 is remarkably
close to that of Sphenobaiera schenckii from the same TC (illustrated here
under Sphenobaiera on p. 222 and previously described by And. & And.
1989, p. 142). This provides a particularly fine example of cuticular corre-
spondence between female fruit and leaf.
H. baccata
tfs 3,4
PRE/F/5943
prep. no. 1003
HAMSHAWVIALES
Hamshawvia
212
d^TRELITZIA 15 (2003)
assemblages
(taphocoenosis)
Sphenobaiera
O Hamshawvia
H. baccata
H. longiped.
H. octosemina
H. linisemina
H. spp. indet
Habitat
Umk 111 Die 2spp
5 7
6 -il'
D1
Lit 111 Dic/Hei
1 2
2 1 -1 -1 -i
Hla 213 Dicelo
49, -
D2
" 212 Die 3spp
ZL -
1 1 1 1
n „
7 -
.1 .1 j j .
Maz 111 Die era
12 -
-1 -1 -1 -1 -
” 211 Hei/Dic
1
- 1 -1 -1 -1
Mat 111 Die dub
18 -
-] -j
Kap 111 Dic/Ris
20 -
_i j j j
Cal 111 Dic/Sph
44 -
D3
Kom 111 Sph/Dic
60 -
-1 -1 -1 -1 -
Vin 111 Die odo
4 [ -
-1 -I -1 -1
Ela 111 " ”
1 -
-! -! -! -
Kra 111 ” ”
i ! -
J j j .1
Kon 223 " "
i ' -
-1 .1 .1 j .
” 222 " ”
io ! -
-1 -1 -1 -1 .
" 111 ” ”
5 -
.
Pen 321 Dic/Ris
5 -
-j -[ -j -]
" 421 Die odo
4 -
- 1 -] -J -]
Nuw 111 Die zub
30
-' -1 -' -
" 211 Die 2spp
3: -
-! -! -! -! -
Qua 111 " ”
20 -
Mak 111 ” ”
i : -
-! -! -! -I ■
Moo 111 Die zub
5 -
_i J j J
Mng 111 Die 2spp
6 -
Qac 111 Hei/Dic
8: -
Bir 211 Sph 2spp
95 -
1 1
“l “I “1 _i
" 311 Hei/Sph
40 -
- -! -j -j
” 111 Sph 2spp
85 -
_ 1 j j j
S
Mol 111 Sph pon
99 -
-i -i -1 -i
Tin 121 Sph 2spp
95 -
-1 -1 -1 -1
Wal 111 Die odo
3 -
Kle 211 Sph pon
100 -
j j j j
Kul 111
99 -
Aas411 Dic/Sph
30 14
-i 1 3 -1 -i 1
" 511 Dicelo
20 -
-! -! -! -!
Gre 111 Sph pon
85 -
> i 1 1
F
Lut 311 " ”
29, 1
j j j y
Kle 111 Hei/Dic
8' -
H
Aas 111 Hei elo
i ! -
’ 311 ” ’’
9 -
-! -! -! -!
Ask 111 Equ sp
1 ! -
1 1 1 1
1 1 1 1
E
Lut 211 ” ”
10 -
• 1 1 1
Total TCs
43j 4
2! 1! 1; 1; 1
Total indivs
% | 24
8 13 l! 1 1
Gondwana Triassic occurrence (elaborated)
Together with Dicroidium and Heidiphyllum , Sphenobaiera is one of
the three clearly dominant gymnosperm foliage genera in the Molteno and
Gondwana Triassic. Its supposed female-fruit affiliate, Hamshawvia ,
remains, however, a rarity and is known outside the Molteno from only a
single specimen in South America and two from Australia.
South America
Frenguelli ( 1942) described Umkomasia cacheutensis, based on a sin-
gle good specimen from the ‘Estratos con Esther ia' in the Cacheuta Fm.,
Upper Triassic of Argentina. HMA has been able to study this specimen in
La Plata (1999) and verify that it is definitely not an Umkomasia. It is
placed here in our new genus as Hamshawvia cacheutensis (Frenguelli
1942) And. & And., comb. nov. This is the only individual known from
South America. Specimens from the Molteno show a similar radiating pat-
tern around the ovules— see PRE/F/21438a, p. 215, tf. 5; pi. 70(7).
Australia
Holmes [1995, pi. 1(1)] illustrated two individuals of a ‘paired ovulate
organ’ from the Nymboida Coal Mine Quarry (see Holmes 2000) of the
Middle Triassic (Ladinian) Basin Creek Fm., Nymboida Coal Measures,
New South Wales. The two clear impressions/compressions (no cuticle)
occur on a single slab along with a number of Sphenobaiera leaves and a
couple of specimens each of Dicroidium and Heidiphyllum. The rarity of
this fruit is confirmed by the fact that these are the only specimens
observed by Holmes (pers. comm.) despite his comprehensive collecting
from the Nymboida locality over many years.
Comparisons beyond Gondwana Triassic
Laurasian Triassic
Hamshawvia has similarities with Leuthardtia and Haitingeria
(both Upper Triassic, Europe) and regarded by Crane (1986) as
probable microsporophylls. However, Meyen (1988) links
Leuthardtia to Stiphorus and considers it female.
Irania hermaphroditica from the Rhaetic of Iran, described by
Schweitzer (1977), has paired megasporophylls and microsporan-
gial strobili attached to an axis. The once-forked axis with paired
megasporophylls is similar to Hamshawvia but differs in the discs
being uniovulate with basal placentation and nonfleshy. The
microsporangia have been compared to those of the extant Ginkgo
biloba and the affiliated leaves are possibly Desmiophyllum armani.
Other ages
Stiphorus is close to Hamshawvia— clearly affiliated with
Glossophyllum— as described by Meyen (in Gomankov & Meyen
1986) from the Tatari na-flota (Late Permian of Eurasia). Stiphorus
is also a strobilus with a once-forked pair of megasporophylls. It dif-
fers in the ovules (seeds) being attached externally and not embed-
ded in the fleshy lamina.
Tab. 47. HamshawvialSphenobaiera, Molteno occurrence;
emphasizing habitat
Habitat
D1 — Dicroidium riparian forest (type 1)
D2 — Dicroidium riparian forest (type 2)
D3 — Dicroidium open woodland
S — Sphenobaiera closed woodland
H — Heidiphyllum thicket
E — Equisetum marsh
? — uncertain o
(O
CO
<
or
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
Hamshawvia
HAMSHAWVIALES
d^TRELITZIA 15 (2003)
213
Evidence for affiliation of organs
The measure of our uncertainty as to which ovulate fruit affiliates with
the diverse and abundant foliage genus Sphenobaiera was first outlined in
some detail in And. & And. (1989, p. 132).
Kindred reinforcement
Meyen (1982 1986, 1987, 1988) reported the forked ovulate genus
Stiphorus (closely comparable to Hamshawvia) as securely affiliating with
Glossophyllum in the Tatarina- flora; and with ‘leafy shoots, Kirjamkenia
lobata , producing both entire (Glossophyllum- like) and palmately dissect-
ed (Sphenobaiera- like) leaves’, from the Permo-Triassic of Siberia. Meyen
(1988) also found Leuthardtia, ‘similar, if not identical' to Stiphorus, thus
securely linking Sphenobaiera-Uke leaves in the Triassic of Switzerland
(the Basel flora of Krausel & Schaarschmidt 1966) with Glossophyllum
leaves from the Donets basin.
Mutual occurrence
Hamshawvia is infrequent, occurring in only four Molteno TCs, but
each of these does yield Sphenobaiera, even if not in high abundance, i.e.
Aas 411 with 30%, Umk 111 5%, Lit 111 1% and Lut 111 less than 1%.
Ginkgoites is a less likely contender as it does not occur at Umk 111 and
Lut 111, while at Aas 411 and Lit 111 it is below 1%. The possibility of
Avatia instead of Hamshawvia affiliating with Sphenobaiera is also less
likely, as it does not occur at Umk 1 1 1 and Lit 111. However, Avatia is very
common (over 50 individuals) at Aas 411 and Bir 111, which both yield
high Sphenobaiera percentages and low Ginkgoites percentages. Fanero-
theca, as we originally considered, could also be a contender for affiliation
with Sphenobaiera on the sole basis of recorded Molteno co-occurrences
(Tab. 48).
Holmes (1995) illustrated two clearly preserved, paired ovulate
organs, almost certainly identifiable as Hamshawvia, from the Middle
Triassic Nymboida Coal Measures, Australia. These appeared on a bed-
ding-plane along with a number of Sphenobaiera leaves.
Organic attachment
A single specimen (Aas 411, PRE/F/12823a,b; pi. 71) from the Molteno
shows four Sphenobaiera leaves and a bifurcating fructification clearly
attached terminally to a short shoot. The forked axis of the organ is distinct
but the two megasporophyll heads are small and not well preserved. We
consider them to be immature fruits. The heads as preserved are one half
the size of those of Avatia and one third the size of Hamshawvia. We accept
that this attached fruit could possibly be Avatia but more likely it is
Hamshawvia. Because of the uncertainty of identification, we have record-
ed this affiliation as Grade 4/5.
Cuticular correspondence
The cuticle of Hamshawvia [tfs 3, 4, p. 211; pis 67(3-9), 68(4—7),
69(4, 5)] from both Lit 111 and Umk 111, is similar to that of Spheno-
baiera, notably in the thickness and quality of preservation, the strongly
papillate cells with gently curved walls, and in the clear ring of lappetate
subsidiary cells. No cuticle is known from Avatia.
Adaptive radiation (Molteno diversity)
Hamshawvia, though both infrequent and rarely preserved, was evi-
dently diverse. Four of the five small Molteno palaeodemes at hand repre-
sent distinct species. These are differentiated by peduncle length and par-
ticularly the number and arrangement of ovules. Two of the species derive
from a single TC (Umk 111), while the others are from very different habi-
tats and stratigraphic levels.
H. baccata—\Jxr\k 111 Die 2 spp (Umkomaas Valley), 6 indivs
Dicroidium riparian forest (mature); Cycle 2b (Indwe Member)
H. longipedunculata— Aas 411 Dic/Sph. (Aasvoelberg), 14 indivs
Sphenobaiera closed woodland; Cycle 1 (Bamboesberg Member)
H. octosemina— Umk 111 Die 2 spp (Umkomaas Valley), 1 indiv.
Dicroidium riparian forest (mature); Cycle 2b (Indwe Member)
H. linisemina— Lut 311 Hei elo (Lutherskop); 1 indiv.
Heidiphyllum thicket; Cycle 3 (Mayaputi Member)
Tab. 48. Hamshawvia! Sphenobaiera , Molteno occurrence;
emphasizing species diversity
HAMSHAWVIALES
Hamshawvia
214
d/TRELITZIA 15 (2003)
Hamshawvia baccata J.M.And. & H.M.And., sp. nov.
Holotype
Specimen'. BP/2/1 124a,b; pis 66(1-4), 67(10, 11).
Assemblage (TC): Umk 111 Die 2spp, Umkomaas Valley.
Preservation: complete strobilus. part and counterpart; compression, in
thinly laminated, carbonaceous (good cuticle), moderately baked, dark
grey shale with good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 6 indivs (3 intact, 3 partial), pis 66, 67.
Sister palaeodemes— 1 (as listed)
Lit 111 Dic/Hei: 2 indivs (2 intact), pi. 68.
Specific diagnosis
A Hamshawvia species with a short axis and megasporophylls with ca
17 ovules radially arranged about a fusiform median vein.
Specific characters
Strobilus: axis short, ca 6 mm to bifurcation.
Megasporophyll: lamina spherical to broadly ovate, with ca 17 ovules
arranged radially about a fusiform vein and connected to simple or
forked vascular traces.
Ovule/seed: elliptical.
Etymology
baccata— bacca (Lat.), berry, berry-like, with reference to the appearance
of the megasporophyll heads.
Comments & comparison
H. baccata is based on two palaeodemes, but particularly that from
Umk 111, representing Dicroidium riparian forest. The diagnostic arrange-
ment and number of ovules seen in the holotype are obscure in the addi-
tional five specimens from Umk 111, which are preserved in varied orien-
tation. However, the additional specimens are similar in size, form and tex-
ture. PRE/F/9877b (tf. 3) shows one megasporophyll in a closed position
folded along the median rib.
Though not much detail can be discerned in the two Lit 111 specimens,
tfs 4, 5 adjacent and pi. 68(1, 2), they are tentatively placed in this species.
They also appear to show ovules with a radiating pattern as in the Umk 111
holotype, and are generally similar in shape and size, and in the form of the
peduncle and pedicel.
Hamshawvia octosemina J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/769; pi. 69(1-7).
Assemblage (TC): Umk 111 Die 2spp, Umkomaas Valley.
Preservation: incomplete megasporophyll, without counterpart; compres-
sion, in thinly laminated, carbonaceous (good cuticle), moderately
baked, dark grey shale with good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimen: 1 indiv (partial); pi. 69(1-7).
Sister palaeodemes— nil.
Specific diagnosis
A Hamshawvia species with megasporophylls bearing 8 ovules (in
rounded-rectangular fields) arranged in rows of 4 on either side of a median
vein.
Specific characters
Strobilus: axis unknown.
Megasporophyll: lamina ovate, with 8 ovules in rounded-rectangular sem-
inal fields, arranged in rows of 4 on either side of a median vein.
Ovule/seed: shape unknown.
Etymology
octosemina— referring to the eight ovules in each megasporophyll.
Comments & comparison
Though based on a single specimen, the diagnostic characters of H. octo-
semina are clear. The species very evidently comes closest to H. linisemina.
Hamshawvia
HAMSHAWVIALES
TRELITZIA 15 (2003)
215
Hamshawvia longipedunculata j.MAnd. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/21778a,b; pi. 70(1-3).
Assemblage (TC): Aas411 Dic/Sph, Aasvoelberg.
Preservation : nearly complete strobilus, part and counterpart; 3D impres-
sion, in thinly laminated, strongly baked, yellowish grey shale with
very good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 13 indivs (2 intact, 11 partial), pi. 70(1-10).
Sister palaeodemes— nil.
Specific diagnosis
A Hamshawvia species with a long axis and megasporophylls with ca
14 ovules in an irregular radial arrangement about a median vein.
Specific characters
Strobilus: axis elongated, ca 25-30 mm to bifurcation.
Megasporophyll: lamina circular, surface wrinkled, with ca 14 ovules in
irregular radial arrangement about an obscure median vein.
Ovule/seed: oval.
Etymology
longipedunculata— referring to the elongated peduncle (actually axis) of
the strobilus.
Comments & comparison
H. longipedunculata differs from H. baccata in the elongate peduncle
and more random arrangement of the ovules in the megasporophyll lami-
na. Specimen PRE/F/21438a, tf. 5, shows best the characteristic radiating
pattern around the ovules found in this species. The feature probably relates
to differential shrinkage between the ovules and surrounding tissue through
dehydration before or during fossilisation.
Hamshawvia linisemina J.M And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/11607; pi. 69(8-10).
Assemblage (TC): Lut 31 1 Hei elo, Lutherskop.
Presen’ation: incomplete strobilus, without counterpart; impression, in
thickly laminated, medium grey shale with moderate cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimen: 1 indiv. (intact), pi. 69(8-10).
Sister palaeodemes— nil.
Specific diagnosis
A Hamshawvia species with an intermediate-length axis and megas-
porophylls with ca 10 ovules (in linear-rectangular fields), arranged in
rows of 5 on either side of a median vein.
Specific characters
Strobilis: axis intermediate, ca 13 mm to bifurcation.
Megasporophyll: lamina ovate, with ca 10 ovules in linear-rectangular
seminal fields arranged in rows of 5 on either side of a median vein.
Ovule/seed: shape unknown.
Etymology
linisemina— referring to the linear-rectangular seminal fields.
Comments & comparison
The Lutherskop (Lut 311) palaeodeme on which this species is based
consists of a single imperfect specimen. Even so, following our taxonomic
guidelines, we feel its identification justified. The specimen falls readily in
the genus Hamshawvia and is most like the single specimen of H. octosem-
ina from Umk 111. The distinguishing feature of H. linisemina is the lin-
ear-rectangular—in contrast to roundly rectangular— areas demarcating the
seeds. This feature is not as clear as in the Umk 111 specimen and could be
misinterpreted. In support of recognising a separate species is the differing
habitat: Umk 1 1 1 has been interpreted as representing Dicroidium riparian
forest, and Lut 3 1 1 Heidiphyllum thicket. Should this be correct, it is more
likely that we are dealing with two species than one.
H. linisemina
HAMSHAWVIALES
Hamshawvia
216
d/TRELITZIA 15 (2003)
Stachyopitys A.Schenck 1867
Type species
Stachyopitys preslii A.Schenck 1867.
Strullendorf, near Bamberg, Bavaria; Rhaetic, Triassic.
Generic concept
A ginkgoopsid male strobilus of lax spicate to bushy form, with sim-
ple to branched microsporophylls bearing terminal spherical heads of radi-
ating microsporangia.
Generic characters (Molteno Fm.)
Attachment: strobilus borne singly on a bulbous short shoot which
dehisces as a unit.
Strobilus: simple, lax, spicate to bushy, radially symmetrical, small to large
(25->150 mm); axis gracile, gently curving to sinuous; microsporo-
phylls numerous, irregularly helical.
Microsporophyll: simple to irregularly branched and bushy, radially sym-
metrical; peduncle gracile; fertile heads one to several, pedicelate;
microsporangia in lax to dense fascicles, few to numerous per head.
Microsporangium: irregularly elliptical to spathulate (0.5-3 mm long);
ornamentation fine, linear, sinuous, forking and converging.
Etymology
Stachyopitys— stachys (Gr.), spike; pitys (Gr.), a pine.
Global range: numerous spp., Pangaea, Tr.-K.
Gondwana Triassic occurrence
S Am— Argentina, 2 TCs (8 indivs).
SAf— Karoo Basin, 27 TCs (539 indivs).
Aus— Queensland, Victoria, N.Z., 7 TCs (10 indivs).
Molteno occurrence
Frequency (F): 27 TCs (of 100 sampled in Molteno).
Diversity (D): 6 species.
Abundance (A): 539 individuals, rare to very rare in top 8 TCs.
Kon 222 Die odo: 35 indivs in 40 man-hrs cleaving ( 9 per 1 man-day) rare
Mat 111 Die dub: 31
” ” 65
” ( 5 ” 1
” ) ”
Cyp 1 11 Die era: 44
” ” 100
” 05 ” 1
” ) ”
Tel 111 Heielo: 45
” ” 90
” ( 4 ” 1
” ) ”
Wal 1 1 1 Die odo: 21
” ” 50
” ( 4 ” 1
” ) ”
Maz 21 1 Hei/Dic: 26
” ” 85
” ( 4 ” 1
” ) ”
Bir 111 Sph 2spp: 113
” ” 550
” (>2 ” 1
” ) ”
Aas 411 Dic/Sph: 67
” ” 512
” (>1 ” 1
” ) v. rare
Though never common, Stachyopitys is the most frequently occurring
gymnospermous microsporangiate genus in the Molteno. Its affiliate,
Sphenobaiera, is the third most prominent foliage genus in the Gondwana
Triassic.
Affiliated organs
Female strobilus: Hamshawvia— Grade 4 (Mut. occ.. Kin. reinf.).
Foliage: Sphenobaiera— Grade 5 (Org. att. ).
Classification & comparison
Suprageneric classification (Hamshawviaceae/Hamshawviales)
From the evidence of the single Molteno specimen (p. 220, tf. 10a, b)
showing the attachment of Stachyopitys (male) and Sphenobaiera (foliage)
to a common base, together with the well established affiliations of both to
Hamshawvia (female), we have an unusually soundly based whole-plant
genus. Following the arguments outlined for Hamshawvia (p. 210), this
combined taxon is placed in the family Hamshawviaceae and order
Hamshawviales within the class Ginkgoopsida.
Intergeneric comparison (Gondwana Triassic)
Stachyopitys is closest to the microsporangiate genera Antevsia and
Pteruchus (particularly P. africanus). While complete, well-preserved
specimens of these three genera can be readily distinguished, fragmentary
or poorly preserved individuals may be confused. Antevsia (from the
Molteno) differs in bearing its relatively few microsporangia in sessile lat-
eral fascicles on projecting flanges along the microsporophylls, and
Pteruchus in its more numerous microsporangia borne abaxially on lami-
nate heads.
Reconstructions
The Stachyopitys strobilus, based on many well-preserved Molteno
palaeodemes, is clearly spicate rather than planate in form. The remarkable
bulbous base, found in many specimens through several palaeodemes
(notably Bir 111, Wal 111, Maz 111, Maz 211), appears to represent a kind
of specialised short shoot. We have made a reconstruction of the single
specimen (PRE/F/4000, p. 211, tf. 2) in which a Stachyopitys ( S . lacrispo-
rangia) strobilus and a Sphenobaiera ( S . sectina) leaf are found mutually
attached to a bulbous base [pi. 81(1-6)]. Reconstructions are also provided
for three of the six Molteno Stachyopitys species.
Stachyopitys
HAMSHAWVIALES
d^TRELITZIA 15 (2003)
217
Gondwana Triassic occurrence (elaborated)
Numerous supposed Stachyopitys specimens have been recorded from
Gondwana (Tab. 49), but some are here placed in Pteruchus (Tab. 64).
Only S. simmondsii from Queensland, based on a complete strobilus, is
accepted by us as a valid species. The remainder, based only on fragmentary
strobili, are regarded as indeterminate at species level. S. simmondsii is
closest to 5. maziramus in size and the attachment to a bulbous base, but
differs in the simple (unbranched) axis and a greater number of sporangia
per head.
Adaptive radiation (Molteno diversity)
Though we have differentiated six species of Stachyopitys , they are
less readily defined than in the putative female (Hamshawvia, four species)
and foliage ( Sphenobaiera , nine species) affiliates. The genus is both
frequent, occurring in 27 TCs, and relatively common, with some 500
curated specimens, but morphologically it is notably conservative. The
diagnostic characters lie chiefly in the shape and size of the strobilus,
together with the shape, size and number of microsporangia per
microsporophyll head. The species showing the greatest frequency is S.
lacrisporangia , which occurs in 23 TCs.
The species derive equally from Dicroidium- and Sphenobaiera-
dominated habitats and from various stratigraphic levels:
S. malatilongus— Mat 111 Die dub (Matatiele), 15 indivs
Dicroidium riparian forest (immature); Cycle 2b (Indwe Member)
S. matatiramus —Mat 111 Die dub (Matatiele), 16 indivs
Dicroidium riparian forest (immature); Cycle 2b (Indwe Member)
S. gypsianthus— Aas 411 Dic/Sph (Aasvoelberg), 44 indivs
Sphenobaiera closed woodland; Cycle 1 (Bamboesberg Member)
S. lacrisporangia— Bir 111 Sph 2spp (Birds River), 57 indivs
Sphenobaiera closed woodland; Cycle 5 (Tsomo Member)
S. rotundisporangia— Bir 111 Sph 2spp, (Birds River), 3 indivs
Sphenobaiera closed woodland; Cycle 5 (Tsomo Member)
S. maziramus — Maz 211 Hei/Dic (Mazenod), 1 indiv.
Dicroidium riparian forest (immature); Cycle 2c (Indwe Member)
Tab. 49.
STACHYOPITYS HYPODIGM, Gondwana Triassic occurrence
Species
Intact-
ness
Molteno
Q.
a
V )
GO
S. matatilongus
S. matatiramus
S. gypsianthus
S. lacrisporangia
S. rotundisporangia
S. maziramus
Intact strobili
Fragmentary "
Isolated megasp.
AUTHOR
SUBREGION
FORMATION
LOCALITY
NAME
Indivs ILLUSTRATION
SOUTH AMERICA
1
1
I 1 I 1 1
1 1 1 1 1
; |
1944c Frenguelli
Cacheuta NA4
24 Potrerillos
YPF old admin, bid.
Stachyopitys anthoides
6 pi 1(1-5), 2(1-2), f 1
- | - [ * ] - 1 - j -
6
-33
1967 Jain & Del.
" ; ”
Minas de Petrolea
"
2 pi 90(16), pi 96(12-13)
- 1 - ! - 1 - 1 - 1 -
2
- 1 1
SOUTH AFRICA
;
!
1 1 1 1 1
1903 Seward
Matatiele KA7
24 Molteno
Kenigha River
Stachyopitys sp.
1 pi 9(2)
_ . . I _ 1 . j . 1 .
1
- i 1 i -
1978-1999 Anderson & Anderson Molteno literature not included in this table
NEW ZEALAND
1 1 1 1 1
1981b Retallack
Benmore Dam NZ4
21 Long Gully
Long Gully
Townrovia petasata
1 pi 11(E), f 3(C)
1
- 1 1 -
1985
Southland ! NZ6
? 1
Mataura Is F47/f7572
Antevsia sp.
1 f 7(7)
- 1 - ! - 1 - 1 - | -
1
- 1 -
AUSTRALIA
; ; j ; ;
1 ;
1898 Shirley
Ipswich/Esk. CM5
24 Blackstone Stage
Denmark Hill (loc 39)
Stachyopitys simmondsi
1 pi 18(2)
1
1 1 - 1 -
1927 Chapman
Victoria Vi2
22 ?
Bald Hill
" cf. annularioides
1 pi 11(25)
- 1 - 1 - 1 - 1 - 1 -
1
- ; - i
1917 Walkom
Ipswich/Esk. CM5
24 Blackstone Stage
Nundah
” annularioides
1 pi 4(6)
- 1 - 1 - 1 - 1 - i -
1
- i -
1962 Townrow (s
oecimen repeated from Shirley 1898)
" simmondsi
* pi 24(2)
*
* | - ; -
1965 !
Hamilton ' Ta?
? T ?
Langloh Coal Mine
Pteruchus petasatus
6 pi 1(B), 14(A-F)
6
12 3
" Hill ef a/.
Brisbane CM6
231 Tingalpa
Petrie's Quarry
” simmondsi
1 : pi T6(5)
1
T - -
GONDWANA TRIASSIC, ‘GEOSTRAT DISTRIBUTION
HAMSHAWVIALES
Stachyopitys
218
d^TRELITZIA 15 (2003)
Evidence for affiliation of organs
Organic attachment
The unique specimen (PRE/F/4000, pi. 81) from
Maz 1 1 1 Die era shows the lower portion of a Spheno-
baiera leaf and part of a Stachyopitys strobilus clearly
attached to a common bulbous base and is thus given a
Grade 5 affiliation. The leaf is close to Sphenobaiera
africana (for further leaves from Maz 111, see And. &
And. 1989; pis 97, 98) and the strobilus to Stachyopitys
lacrisporangia.
The only other known specimen involving related
taxa in apparent organic connection was described by
Yang (1986) from China (Xujanhe Fm. in Xing-Fong,
Wan-Gu region of Dazu, Sichuan). The male strobilus
was close to Stachyopitys and has been named Spheno-
baieroanthus sinensis. The exact nature of attachment
is difficult to discern from the photographs, but cer-
tainly there is a clear shoot with numerous Spheno-
baierocladus sinensis leaves attached.
Mutual occurrence
The most convincing instance of co-occurrence is
at Bir 111 where Sphenobaiera comprises 85% of the
assemblage and Stachyopitys is represented by 1 1 3 cat-
alogued specimens (Tab. 50). Similar good associa-
tions are recorded from Aas 411 and Umk 111. Some
of the Stachyopitys species are possibly affiliated with
Fanerotheca which also occurs in high numbers at Bir
111, Aas 4 1 1 and Umk 111 (see also Wal 111, Kap 111,
Lut 111). For further discussion consult affiliation text
under Fanerotheca. Another uncertainty is that there
are six TCs where Stachyopitys is present but Spheno-
baiera is absent, i.e. Cyp 111, Tel 1 1 1 , San 111, Kra 311,
Mor 1 1 1, Kan 112. Fanerotheca appears at the first three
of these TCs.
Kindred reinforcement
Associations of Sphenobaiera and Stachyopitys
were recorded from the Basel flora, Switzerland
(Keuper, Upper Triassic), by Leuthardt (1903). From
the Early Fiassic of Bayreuth, Germany, Kirchner &
Van Konijnenburg-Van Cittert (1994) described the
female organ Schmeissneria microstachys attached as
a fascicle with numerous simple elongate leaves to a
short shoot. This unique genus is considered by them
to be of ginkgoalean affinity. They regarded the affili-
ated male as Stachyopitys preslii (not yet found
attached). It was conceded that "there may be other
plants within the Ginkgoales carrying Stachyopitys
fructifications’.
Tab. 50. Stachyopitys, Molteno occurrence
Stachyopitys
HAMSHAWVIALES
rV TRELITZIA 15 (2003)
219
Stachyopitys matatilongus J.M.And. & H.M.And., sp. nov.
Holotype
Specimen'. PRE/F/9250a,b; pis 72(1), 73(1-3).
Assemblage (TC): Mat 1 1 1 Die dub, Matatiele.
Preservation', intact strobilus, tip and base missing, part and counterpart; im-
pression, in thickly laminated, olive-grey shale with moderate cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 15 indivs (6 intact, 8 partial, 1 isolated), pis 72(1-4), 73(1-6).
Sister palaeodemes— 1 only (as listed)
San 111 Die era: 1 indiv. (intact).
Specific diagnosis
A Stachyopitys species with long, linear strobili, bearing simple to multi-
branched microsporophylls with narrowly elliptical microsporangia.
Diagnostic characters
Attachment: unknown.
Strobilus: linear, long (up to >140 X 20 mm).
Microsporophylh peduncles simple to forked to multiple-branched.
Fertile head: ultimate heads with 3 or 4 microsporangia.
Microsporangium: narrowly sinuously elliptical.
Etymology
matatilongus— ref erring to the long strobilus from the Matatiele locality.
Comment & comparison
With its long spicate strobili, S. matatilongus is by far the most impres-
sive and distinctive of the Stachyopitys species.
Stachyopitys matatiramus J.M.And. & H.M.And., sp. nov.
Holotype
Specimen BP/2/31 15; pi. 74(5, 7).
Assemblage (TC): Mat 111 Die dub, Matatiele.
Preservation: intact strobilus, base missing, no counterpart; impression, in
thickly laminated, olive-grey shale with moderate cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 16 indivs (11 intact, 5 partial), pi. 74(1-13).
Sister palaeodemes— 1 only (as listed)
Tel 111 Hei elo: 1 indiv. (partial).
Specific diagnosis
A Stachyopitys species with small compound (fasciculate) strobili,
bearing simple microsporophylls with narrowly elliptical microsporangia.
Diagnostic characters
Attachment: with up to 4 strobili arising from a linear short shoot.
Strobilus: linear, spicate. relatively small (to 40 X 7 mm).
Microsporophylh peduncles almost exclusively simple.
Fertile head: with ca 4 to 7 microsporangia.
Microsporangium: narrowly sinuously elliptical.
Etymology
matatiramus— matati, for the type locality; ramus (Lat.), branch, referring
to the branched structure.
Comment & comparison
S. matatiramus occurs in the same TC (Mat 111) as S. matatilongus
and is distinguished by its far smaller size and the branched structure of the
strobilus. The species is reminiscent of Antevsia mazenodensis (affiliate of
Peltaspermum) which, however, has sessile microsporangial clusters (see
p. 155).
HAMSHAWVIALES
Stachyopitys
220
c/tRELITZIA 15 (2003)
Stachyopitys gypsianthus J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/1 3074a, b; pi. 75(1, 2, 8).
Assemblage (TC): Aas 411 Dic/Sph, Aasvoelberg.
Preservation: complete strobilus, part and counterpart; impression, in thinly
laminated, strongly baked, yellowish grey shale with very good cleavage.
Reference palaeodeme
Assemblage (TC): as for the holotype.
Specimens: 44 indivs (21 intact, 23 fragm.), pi. 75(1-9).
Sister palaeodeme— 2 (best 1 listed)
Lut 111 Hei/Dic: 10 indivs (1 intact, 8 partial, 1 isolated).
Specific diagnosis
A Stachyopitys species with medium-sized linear strobili, bearing sim-
ply branched microsporophylls with narrowly elliptical microsporangia.
Specific characters
Attachment: unknown.
Strobilus: linear, spicate, of medium size (up to ca 60 X 12 mm).
Microsporopliyll: peduncles branching distally.
Fertile head: ultimate heads with ca 3-6 microsporangia.
Microsporangium: narrowly sinuously elliptical.
Etymology
gypsianthus— gyps (Gr.), vulture, with reference to the type locality, Aasvoel-
berg (Afrikaans), meaning "vulture mountain’; anthos , flower.
Comment & comparison
S. gypsianthus is most like 5. matatilongus from Mat 111, but differs
in the strobili being consistently shorter, smaller and the pedicels not hav-
ing multiple branching. It differs from the somewhat similar S. matatira-
mus in bearing simply branched microsporophylls.
Stachyopitys lacrisporangia J.M.And. & H.M.And., sp.
nov.
Holotype
Specimen: BP/2/5073a,b; pi. 77(1-5).
Assemblage (TC): Bir 111 Sph 2spp, Birds River.
Preservation: complete strobilus attached to bulbous base, part and coun-
terpart; impression, in thinly laminated, yellowish grey shale with very
good cleavage.
Reference palaeodeme
Assemblage (TC): as for the holotype.
Specimens: robust group— 53 indivs (38 intact, 1 1 partial, 4 isolated),
slender group— 57 indivs (44 intact, 10 fragm., 3 isolated), pis 76, 77.
Sister palaeodemes— 22 (best 2 listed)
Tel 111 Hei elo: 44 indivs (21 intact, 16 fragm., 7 isolated), pi. 78(1-12).
Wal 111 Die odo: 20 indivs (8 intact, 8 partial, 4 isolated), pi. 78( 1 — 4).
Specific diagnosis
A Stachyopitys species with short, linear to broadly spicate strobili,
bearing simple microsporophylls with clavate or tear-shaped microsporan-
gia.
Specific characters (based on reference palaeodeme, Bir 111)
Attachment: with a single strobilus attached to a bulbous short shoot.
Strobilus: linearly to broadly spicate, relatively small (up to ca 40 X 12 mm).
Microsporophyll: peduncles simple.
Fertile head: with numerous (up to >20) microsporangia.
Microsporangium: clavate or tear-shaped.
Etymology
lacrisporangia— lacrima (Lat.), tear, referring to the tear-shaped microspo-
rangia.
Comment & comparison
This is the most frequent and abundant of the Stachyopitys species
occurring in the Molteno. The many palaeodemes show a wide range of
variation, but at present we find no justification for distinguishing further
species within the group. Two forms with the robust and the slender axes are
separated out in Tab. 50. At Bir 111, about equal numbers of specimens occur
in these two groups. The holotype derives from the more robust category.
The attached specimen from Maz 111 (tf. 10 adjacent) is regarded as
belonging to S. lacrisporangia.
Stachyopitys
HAMSHAWVIALES
c/tRELITZIA 15 (2003)
221
Stachyopitys rotundisporangia J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: BP/2/5072; pi. 79(5-9).
Assemblage (TC): Bir 111 Sph 2spp, Birds River.
Preservation: complete strobilus attached to bulbous base, without counter-
part; impression, in thinly laminated, yellowish grey shale with very
good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 3 indivs (2 intact, 1 fragm.),'pl. 79(5-9).
Sister palaeodemes— 1 only (as listed)
Aas 41 1 Dic/Sph; 7 indivs (3 intact, 3 partial, 1 isolated).
Specific diagnosis
A Stachyopitys species with small, broadly spicate strobili, bearing
simply branched microsporophylls with spherical microsporangia.
Specific characters (based on reference palaeodeme, Bir 111).
Attachment: with a single strobilus attached to a bulbous short shoot.
Strobilus: broadly spicate, small (up to call X 11 mm).
Microsporophyll: peduncles branching distally.
Fertile head: ultimate heads with ca 3 or 4 microsporangia.
Microsporangium: spherical.
Etymology
rotundisporangia— rotundus (Lat.), almost circular, rounded, referring to the
spherical microsporangia.
Comment & comparison
The reference palaeodeme of this species is represented by the holotype
(a clearly preserved complete specimen) and only two further individuals.
It differs from other Stachyopitys species in the small spherical microspo-
rangia.
Stachyopitys maziramus J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/4652; pi. 80(1, 2).
Assemblage (TC): Maz 21 1 Hei/Dic, Mazenod.
Presen’ation: complete strobilus attached to bulbous base, without coun-
terpart; compression, in thinly laminated, carbonaceous (poor cuticle),
medium grey shale with moderate cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimen: holotype only.
Sister palaeodemes— nil.
Specific diagnosis
A Stachyopitys species with short compound (fasciculate) strobili, bearing
multibranched microsporophylls with ovate-elliptical microsporangia.
Specific characters (based on reference palaeodeme. Maz 211).
Attachment: with 2 or more strobili arising from a bulbous short shoot.
Strobilus: short, fasciculate (ca 12 X 12 mm).
Microsporophyll: peduncles variously multibranched.
Fertile head: ultimate heads with many (up to >15) microsporangia.
Microsporangium: ovate-elliptical.
Etymology
maziramus— mazi. for the type locality, Mazenod; ramus (Lat.), branch, refer-
ring to the branched structure of the strobilus.
Comment & comparison
S. maziramus , represented only by the holotype, is the rarest member
of the genus. It differs from other Stachyopitys species in its compound
multibranched axes. The 25 additional specimens of the genus from the
Maz 211 TC are clearly different and fall in the robust form of S. lacri-
sporangia.
HAMSHAWVIALES
Stachyopitys
222
(S^' TRELITZIA 15 (2003)
Sphenobaiera Florin 1936
Type species
Sphenobaiera spectabilis (Nath. 1906) Florin 1936.
Stabbarp, Scania, Sweden; Liassic, Jurassic.
Generic concept
A ginkgoopsid leaf with indistinct petiole, deeply dichotomously
dividing wedge-shaped lamina and forking subparallel venation.
Generic characters (Molteno Fm.)
Attachment : leaves fascicled or in a close spiral at the end of short shoots.
Leaf, linear to broadly wedge-shaped with margins diverging at <90°, lam-
ina simple to deeply dichotomously dividing, with each segment to
several times forked; petiole not distinct; veins subparallel, repeatedly
forking, relatively well spaced, converging distally to end separately at
segment apex.
Cuticle : see And. & And. ( 1989. p. 133); this vol., tfs 1-4 below.
Etymology
Sphenobaiera— spheno (Gr.) wedge; Baiera, a genus for ginkgoopsid leaves.
Global range: numerous species, Pangaea, P.-U.K.
Gondwana Triassic occurrence (after And. & And. 1989)
Frequency (F): 26 degree squares (of the 84 across Gondwana).
Ubiquity (U): 5 continents (of 5 comprising Gondwana).
Diversity (D): 12 foliage species.
Abundance (A): 30% (the norm in Molteno TCs).
Longevity (L): 26 myrs (Spathian to later Norian).
Colonisation success: FUDAL rating 26/5/12/30/26 = 99.
High success (Grade 4); Sphenobaiera was the third most prominent
genus in the Gondwana Triassic; it was frequent, ubiquitous, diverse,
abundant and long-lived.
Endemism: of the 12 Gondwana Triassic species, six occur more or less
widely through the realm, one is a basin endemic and five are single-
assemblage endemics.
Molteno occurrence
Frequency (F): 43 TCs (of 100 sampled in the Molteno).
Diversity (D): 9 species.
Abundance (A): monodominant (>70%) in 7 TCs; co-dominant (20-69%)
in 7 TCs; abundant to occasional ( 1-19%) in 9 TCs; rare to very rare
(<1%) in 20 TCs.
Habit: probably woody shrubs to large trees.
Preferred habitat: Sphenobaiera is the monodominant or co-dominant taxon
in lake deposit TCs. It is relatively rare elsewhere.
Affiliated organs
Female strobilus: Hamshawvia— Grade 4/5 (Org. att.. Cut. cor., Mut. occ.,
Kin. reinf.).
Male strobilus: Stachyopitys— Grade 5 (Org. att., Mut. occ.).
See relevant text under Hamshawvia and Stachyopitys. Both these gen-
era have been found in organic connection with Sphenobaiera leaves, but
for reasons previously outlined, a Grade 4/5 affiliation is given for
Hamshawvia. Slender stems with irregularly spiralled leafless short shoots
(much like in extant Ginkgo) occur in some Sphenobaiera woodland TCs
(e.g. Bir 111 and Aas 411). It is likely that these originally bore Spheno-
baiera foliage.
2
4
guard cells
x500
Classification & comparison
Intergeneric comparisons
Gondwana Triassic— In gross leaf morphology, notably the dicho-
tomising nature of both lamina and venation, Sphenobaiera and Ginkgoites
are clearly alike. Certain linear-leaved forms of Sphenobaiera and Dicroi-
dium (i.e. S. pontifolia and D. elongatum subsp. argentinum ) do, however,
prove virtually indistinguishable other than through examination of their
cuticles.
Other ginkgoalean genera— The twelve species of Sphenobaiera from
the Gondwana Triassic conform in the main to the concept of the genus, but
in several instances and in certain features, transgress (or approach very
closely) the boundaries of adjacent ginkgoalean or czekanowskialean genera.
Such genera include Baiera , Pseudotorellia, Eretmophyllum , Czekanowskia
and Phoenicopsis.
Other genera— The lanceolate undivided forms of Sphenobaiera ( S .
insecta and S. calensis) are superficially very like the Gondwana Triassic
conifer Heidiphyllum, but are readily separated on the basis of cuticle and
the absence of interveinal striae.
Interspecific comparisons
As is generally the case, diversity in the Molteno is more readily
recognised in the foliage (Sphenobaiera. nine species) than in the female
(Hamshawvia, four species) and male (Stachyopitys, six species) affiliates.
Many of the Molteno palaeodemes of Sphenobaiera, as in Dicroidium, are
complex and bridge two or more of the defined species. Again, like
Dicroidium, Sphenobaiera actively colonised and diversified through much
of the Triassic, with hybridisation and polyploidy probably widespread—
suggesting a reticulate rather than punctuate evolutionary model. For a full
systematic coverage of the nine Molteno species, with comprehensive pho-
tographic cover and line drawings of palaeodemes, refer to And. & And.
(1989, pp. 130-217).
Sphenobaiera
HAMSHAWVIALES
r/TRELITZIA 15 (2003)
223
HAMSHAWVIALES
Sphenobaiera
dfi. TRELITZIA 15 (2003)
Holotype
Umkomaas Valley
(Umk 1 1 1 Die 2spp)
bp/2/1 124a
BP/2/ 11 24a h/
BSS8
BP/2/1 124a
; PRE/F/9877b
| PRE/F/9877b®g
PRE/F/18731
PRE/F/18731
PRE/F/771b
PRE/F/771b v
m
g'gOT'v;. nxi/m arsF.,
-
4
Jfcr
Jfjji
E®/v
Hamshawvia baccata
pi. 66
HAMSHAWVIALES
225
^/tRELITZIA 15 (2003)
HAMSHAWVIALES
pi. 67
Hamshawvia baccata
226
^VtRELITZIA 15 (2003)
PRE/F/5943b
iSjO BP/2/1741 '(«
BP/2/1741
Little Switzerland
(Lit 111 Dic/Hei)
PRE/F/5943
PRE/F/5943
PRE/F/5943
Hamshawvia baccata
pi. 68
HAMSHAWVIALES
227
d^TRELITZI A 15 (2003)
X100
Umkomaas Valley
(Umk 1 1 1 Die 2spp)
H. octosemina
all PRE/F/769 Holotype
PRE/F/11607
Holotype
Lutherskop
(Lut 3 1 1 Hei elo)
H. linisemina
jj PRE/F/11607
PRE/F/11607
HAMSHAWVIALES
pi. 69
Hamshawvia spp,
& : TRELITZIA 15 (2003)
PRE/F/21778a
Holotype
PRE/F/21188
Aasvoelberg
(Aas 411 Dic/Sph)
PRE/F/12932b
3 i PRE/F/12932b
■ PRE/F/21778a
PRE/F/21778a
PRE/F/21779
PRE/F/21779
| PRE/F/2 1438b
'RE/F/21438a
Hamsliawvia longipedunculata
pi. 70
HAMSHAWVIALES
TRELITZIA 15 (2003)
229
PRE/F/12823b
PRE/F/12823b
Aasvoelberg
(Aas 41 1 Dic/Sph)
fpRE/F/12823a
HAMSHAWVIALES
pi. 71
Hamshawvial Sphenobaiera
PRE/F/9248
asaflfct. r?-~
PRE/F/9247b _
PRE/F/9250b
Holotype
PRE/F/9246
Stachyopitys matatilongus
pi. 72
HAMSHAWVIALES
230
Matatiele
(Mat 1 1 1 Die dub)
c/tRELITZIA 15 (2003)
231
TRELITZIA 15 (2003)
Matatiele
(Mat 111 Die dub)
PRE/F/9250a
PRE/F/9250b
X5
>6r !
PRE/F/9242a
PRE/F/9242a i
HAMSHAWVIALES
pi. 73
Stachyopitys matatilongus
d^TRELITZIA 15 (2003)
7 )& '
PRE/F/9245 x2
PRE/F/9238 x2
BP/2/3115
Holotype
PRE/F/9244 x2
PRE/F/9243 x2
Matatiele
(Mat 1 1 1 Die dub)
PRE/F/1979
PRE/F/1934
BP/2/3115
‘■"iK’/f1 ''I**' .
PRE/F/9238
PRE/F/9244
PRE/F/9238
f PRE/F/9244
PRE/F/9244
pi. 74
Stachyopitys matatiramus
HAMSHAWVIALES
233
d^TRELITZIA 15 (2003)
PRE/F/21081a X2
PRE/F/13074a
^ PRE/F/20658a
PRE/F/21081b
PRE/E/2 1789a
PRE/F/13074a
Holotype
Aasvoelberg
(Aas 411 Dic/Sph)
PRE/F/21081a
\ PRE/F/20658a
PRE/F/ 13074a
HAMSHAWYIALES
pi. 75
Stachyopitys gypsianthus
234
d?TRELITZIA 15 (2003)
BP/2/5070a
BP/2/5070a X2
Birds River
(Bir 111 Sph 2spp)
PRE/F/10060
- ' ' ' 1
BP/2/5070a
X40
* PRE/F/10747b - X2
:<>-
PRE/F/10055 if X2 >
BP/2/5076 X2
Stachyopitys lacrisporangia
pi. 76
HAMSHAWVIALES
r/ TRELITZIA 15 (2003)
235
HAMSHAWVIALES
pi. 77
Stachyopitys lacrisporangia
BP/ 2/ 5 626b
PRE/F/7704a
PRE/F/18219 x2
PRE/F/217321b
PRE/F/17320b
fcssssss ^ ■
PRE/F/17321b
Telemachus Spruit
(Tel 1 1 1 Hei elo)
PRE/F/17320b
PRE/F/17321b
PRE/F/17320a
PRE/F/17320b
PRE/F/17321b
Stachyopitys lacrisporangia
pi. 78
HAMSHAWVIALES
237
HAMSHAWVIALES
pi. 79
Stachyopitys spp.
TRELITZIA 15 (2003)
PRE/F/2208b
all BP/2/5072
Holotype
Birds River
(Bir 111 Sph 2spp)
PRE/F/9436a
X2
PRE/F/9445'y'
Waldeck
(Wal 111 Die odo)
PRE/F/9437
S. lacrisporangia
!3S
d?TRELITZIA 15 (2003)
PRE/F/4652
PRE/F/4672 X2
PRE/F/4652
X5 PRE/F/4654b X2
Flolotype
Mazenod
(Maz 211 Hei/Dic)
PRE/F/11719
PRE/F/4666
PRE/F/1 1717
PRE/F/11719
PRE/F/11715
Stachyopitys maziramus
pi. 80
HAMSHAWVIALES
TRELITZIA 15 (2003)
all PRE/F/4000
Mazenod
(Maz 1 1 1 Die era)
all PRE/F/4000
HAMSHAWVIALES
Stachyopitysl Sphenobaiera
240
d^TRELITZIA 15 (2003)
GINKGOOPSIDA S.V.Meyen 1987
UMKOMASIALES S.V.Meyen 1984
UMKOMASIACEAE S.V.Meyen 1984
Umkomasia H.H.Thomas 1933
Synonyms
Pilopliorosperma H.H.Thomas 1933 (see Holmes 1987).
Spermatocodon H.H.Thomas 1933.
Type species
Umkomasia macleanii H.H.Thomas 1933.
Umkomaas Valley, Karoo Basin, S. Africa; Camian, Triassic.
Generic diagnosis emended
A ginkgoopsid strobilus of lax paniculate form with megasporophylls
bearing 1-7 pairs of uni-ovulate cupules and seeds with bifid, curved
micropyles.
Generic characters
Strobilus : simple, lax, paniculate, small to large (length 40->150 mm);
axis generally robust, erect, gradually tapering; megasporophylls several
to many, attachment very variable, from semi-opposite or alternate,
apparently planar to irregularly helical to irregularly decussate.
Megasporophyll : simple, pedunculate, planar to spicate; bracteoles prominent,
singly or in opposite pairs, axillary or along peduncles; ovuliferous
cupules recurved, pedicellate to sessile, in 1-7 opposite to subopposite
pairs.
Cupule : small to large (3-20 mm deep), circular to roundly triangular to
oval, partially flattened, uni-ovulate, fully to partially enclosing the
ovule, generally deeply splitting at maturity into 2 or 4 regular lobes.
Ovule/seed: slightly asymmetrical, size and shape as per cupules, generally
moderately platyspermic and weakly winged; micropyle pronounced,
bifid, outwardly curved.
Etymology
Umkomasia— named by Thomas (1933) after the type locality.
Global range: several spp., Gondwana, Tr. (SCY-CRN)
First: Umkomasia sp. indet. ( Dicroidium 'flowers’). White (1986), Clarence
Siding, Banks Wall Fm., Blue Mts, Australia.
Last: Umkomasia sp. indet. (Baiera tenuifolia ), Johnston (1888), Lord’s Hill,
Brady Fm., Hobart, Tasmania.
Gondwana Triassic occurrence
SAm— N. & S. Argentina, 3TCs (6 indivs).
SAf— Karoo Basin, 22 TCs (503 indivs).
Aus— Eastern Australia, 18 TCs (48 indivs); New Zealand 4 TCs (8 indivs).
Ant— Allan Nunatak, 1 TC (1 indiv.).
Molteno occurrence
Frequency (F): 22 TCs (of 100 sampled in the Molteno).
Diversity (D): 8 species.
Abundance (A): 503 indivs total; rare to very rare in top 8 TCs.
Mat 111 Die dub: >75 indivs in 65 man-hrs (12 per 1 man-day) rare
Kra 1 1 1 Die odo:
14 ”
” 20
( 7 ”
” ) ”
Kon 222 Die odo:
22 ”
” 40
( 6 ”
” ) ”
Hla 213 Die elo:
32 ”
” 60
( 6 ”
” ) ”
Maz 211 Hei/Dic:
46 ”
” 85
( 6 ”
” ) ”
Nuw 111 Die zub:
11 ”
” 21
( 5 ”
” ) ”
Umk 1 1 1 Die 2 spp:
197 ’’
” 400
( 5 ”
” ) ”
Lit 111 Dic/Hei:
51 ”
” 550
( 1 ”
” ) very rare
We include figures above only for those eight TCs yielding the greatest
abundance of Umkomasia.
Affiliated organs
Male strobilus: Pteruchus — Grade 4 (Mut. occ., Cut. corn).
Foliage: Dicroidium— Grade 4 (Mut. occ., Cut. corr.).
Classification & comparison
Suprageneric classification (Umkomasiaceae/Umkomasiales)
Umkomasia , along with its male ( Pteruchus ) and foliage ( Dicroidium )
affiliates— and the related ovulate strobilus Fanerotheca— are recognised
here as the sole representatives of the family Umkomasiaceae and order
Umkomasiales. Peltaspermum and the other five Gondwana Triassic ovulate
genera included in the Ginkgoopsida (Tab. 30) are all considered too remote
morphologically to be included in the same family or order.
Intergeneric comparison (Gondwana Triassic)
Fanerotheca , in its strobilus and cupulate megasporophylls, comes
closest to Umkomasia. It differs mostly in the nature of its strongly winged
seeds. Kannaskoppia resembles Umkomasia in its superficially similar
cupulate units, but differs strongly in its forked architecture. Most other
Molteno ginkgoopsid genera differ in not bearing cupulate structures.
Karibacarpon (not known from the Molteno) from the Upper Triassic
Ripple-marked Flags, Lake Kariba, Zimbabwe, is a somewhat similar genus,
but differs in its cupules splitting asymmetrically close to the pedicel attach-
ment.
Cuticles
Potential sample: Lit 111, 51 indivs; Umk 111, 197 indivs.
Macerated: none in this work; see Thomas ( 1933).
Preservation grade: Grade 4-5.
Diagnostic characters: cells isodiametric to linear-oblong, end walls square
to oblique, walls gently curved to sinuous, nonpapillate to papillate;
trichome bases present; stoma orientation random; subsidiary cells
brachyparacytic, nonlappetate; guard cells narrowly elliptic.
Comment: —
Significance:
Classification— The Umkomasia cuticle as described by Thomas (1933)
and Townrow ( 1 962) is comparable to that of Dicroidium as described by us
(And. & And. 1983, 1989) and matches the general morphology characteris-
ing the class Ginkgoopsida. The main features in common are the narrowly
elliptic guard cells and largely brachyparacytic and nonlappetate subsidiary
cells.
Affiliations— The cuticle of Umkomasia supports affiliation with the
leaf genus Dicroidium as first suggested by Thomas ( 1933). In the features
described above (under ‘Classification’), the Umkomasia cuticle is clearly
more like that of Dicroidium than any of the other Molteno ginkgoopsid
foliage genera, e.g. Lepidopteris, Sphenobaiera, Ginkgoites and Dejerseya.
These latter taxa are all characterised by strongly lappetate, actinocyiic sub-
sidiary cells. The cuticles illustrated by Thomas are, in general, closest to those
of Dicroidium odontopteroides. >
Reconstructions
Strobilus
The well-preserved collection of intact strobili from the Molteno has
added greatly to our knowledge of Umkomasia. The R4 reconstructions
opposite are based on the almost complete strobili of U. macleanii from
Umklll (PRE/F/6602a,b), and of U. quadripartita from Mat 111
(BP/2/9157a,b). Minimal reconstruction was necessary and this mainly by
straightening out the tip and adding the missing megasporophylls. No com-
plete reconstructions for the other Molteno species are provided.
Comparative R3 sketches (p. 244) have been made of the megasporophyll,
cupule and seed for all eight Molteno species.
Megasporophyll
The position and number of bracteoles on the megasporophyll varies
uncertainly with the species. The bracteoles, being small and delicate, may
be poorly preserved and thus may have been overlooked or become
detached. In U. quadripartita there appear two pairs on the megasporophyll
peduncle (at base and midway), with single bracteoles also occurring iso-
lated along the main axis as shown in the reconstruction.
Cupule
The partly decussate nature of the cupules shown in U. quadripartita
is supported by the almost 3D preservation occurring at Aas 411, where U.
decussata has sessile and undoubtedly decussate cupules.
Seeds/ovules
From Umkomasia ovules that are preserved as compressions it is diffi-
cult to determine whether, in life, they were bilaterally or radially symmet-
rical. At Aas 411, where they are preserved in an almost 3D state, the
cupules are flattened even when viewed laterally (pi. 86). If they were radi-
al then this view should also be circular. We interpret Umkomasia
ovules/seeds in general as being bilaterally symmetrical and platyspermic
in cross section.
Umkomasia
UMKOMASIALES
d^TRELITZIA 15 (2003)
241
U. macleanii
based on
PRE/F/6602a,b
pl 82(3,4)
U. monopartita
UMKOMASIALES
Umkomasia
242
d/TRELITZIA 15 (2003)
* unpublished
* specimens repeated — number of individuals therefore not given
Umkomasia
UMKOMASIALES
c/tRELITZIA 15 (2003)
243
Exclusions & synonyms
Gondwana Triassic
A new Umkomasia species, U. unirama , was described from the
Shackleton Glacier region, Antarctica, from either the Upper Fremouw or
Lower Falla Fm. by Axsmith et al. (2000). This unique strobilus has an axis
bearing a whorl of four to eight cupules at the end of elongate stalks
(peduncles). The whorled structure differentiates it from any other known
Umkomasia species. It is somewhat similar to the genus Matatiella (p. 172)
which has an ovule/seed-bearing palmate structure but no clear cupules.
The Antarctic species, with the single axis bearing cupules in a subapical
whorl would considerably expand the generic diagnosis of Umkomasia. We
consider that its placement in Umkomasia is doubtful and that it should
probably be placed in a new genus and possibly a new family and order in
the Ginkgoopsida. We have not included it in our hypodigm for
Umkomasia.
The fertile specimens referred to U. unirama are attached to short
shoots on a branch similar to the mature long shoot in extant Ginkgo bilo-
ba. Another specimen from the same outcrop bears a single Dicroidium leaf
apparently attached directly to a mature long shoot which also has short
shoots similar to those on the fertile specimen. This would be the first
record of Umkomasia attached to a short shoot, and of a Dicroidium leaf
attached singly and laterally to a long shoot and not fascicled and terminal
as suggested by our Molteno material. As the ovulate structure and leaf are
not actually attached to the same branch or the leaf to a short shoot, the
‘unequivocal evidence’ (Axsmith et al. 2000) for Grade 5 affiliation is
missing.
Structurally preserved ovules have been described from the Upper
Fremouw Fm. of Antarctica as Ignotospermum monilii (Perovich & Taylor
1989). These are radially symmetrical and similar in shape and size (3.0 oo
3.75 mm) to seeds/ovules of Umkomasia macleanii but unlike any of the
other Molteno ovulate genera. They have not been found with the typical
bifid micropyle or with the associated cupules as in Umkomasia.
Holmes (1987) synonymised the genera Pilophorosperma and
Karibacarpon with Umkomasia and mentioned that Spermatocodon (based
on one poor specimen by Thomas 1933) was problematical. We consider
Spermatocodon to be a poorly developed fragment of an unidentifiable
Umkomasia species and it is here synonymised with Umkomasia.
The ovulate genus Karibacarpon , described together with its seed
Sengwacarpon by Lacey (1976) from the Upper Triassic Ripple-marked
Flags, Lake Kariba, Zimbabwe, and by Holmes & Ash (1979) from the
Camden Head Claystone of the Lome Basin, eastern Australia, is here
regarded as distinct from Umkomasia. It differs in the cupule splitting
asymmetrically close to the pedicel attachment. However, at both the
Zimbabwean and Australian localities, Karibacarpon is closely associated
with Dicroidium leaves.
Gondwana Triassic occurrence (elaborated)
Umkomasia , after Fanerotheca, is the second most frequent (22 TCs,
Tab. 52) and the most abundant (503 indivs) female strobilus in the
Molteno (Tabs 6b, 9a, 11). It is the most widely recorded through the
Gondwana Triassic. The largest collections are from South Africa (Tab.
52), followed by eastern Australia (Tab. 51).
The first described records of the female fruit of Dicroidium , now
placed in Umkomasia , are those of Johnston (1888) from Tasmania.
Holmes (1987) described three species: U. polycarpa from the Esk Fm. of
Queensland; U. distans and U. sessilis from the Middle Triassic Basin
Creek Fm. of eastern Australia. However, all other specimens (see Tab. 5 1 )
are too inadequately preserved or recorded to identify to species level.
Adaptive radiation (Molteno diversity)
In his original monograph describing the genus Umkomasia, based on
‘about 30 specimens of seed-bearing structures’ from Umkomaas (Umk
111), Thomas (1933) recognised three genera (Umkomasia, Pilophoro-
sperma and Spermatocodon) including 1 1 species. From our more exten-
sive collection of 197 specimens from Umk 1 1 1, we believe that we have a
better understanding of the natural range of variation within the once living
populations. Based on the collections of both Thomas and ourselves, we
recognise two palaeodemes and two species: U. macleanii is by far the
most abundant; U. monopartita (also from Mat 111) is rare. The latter was
not found or described by Thomas (1933).
From our collections— ca 503 individuals from 22 TCs— we recognise
eight species of Umkomasia. The diagnostic characters are based on the
size and laxness or compactness of the strobilus and on the number and
arrangement of cupules in the megasporophyll. This level of diversity is not
reflected in the male affiliate Pteruchus, with only three species recog-
nised.
Most species derive from Dicroidium riparian forest and from Cycle 2
(Indwe Member). The eight species, based mostly on substantial palaeo-
demes from different TCs, are as follows:
U. macleanii— Umk 111 Die 2 spp (Umkomaas Valley), 175 indivs
Dicroidium riparian forest (mature); Cycle 2b (Indwe Member)
U. bracteolata—H\a 213 Die elo (Hlatimbe Valley), 3 indivs
Dicroidium riparian forest (immature); Cycle 2b (Indwe Member)
U. quadripartita—Mm 111 Die dub (Matatiele), 30 indivs
Dicroidium riparian forest (immature); Cycle 2b (Indwe Member)
U. decussata—Aas 411 Dic/Sph (Aasvoelberg), 7 indivs
Sphenobaiera closed woodland; Cycle 1 (Bamboesberg Member)
U. monopartita— Mm 111 Die dub (Matatiele), 45 indivs
Dicroidium riparian forest (immature); Cycle 2 (Indwe Member)
U. gracilliaxis— Lit 111 Dic/Hei (Little Switzerland), 42 indivs
Dicroidium riparian forest (mature); Cycle 2a (Indwe Member)
U. cupulata— Kon 222 Die odo (Konings Kroon), 20 indivs
Dicroidium open woodland (floodplain); Cycle 2f (Mayaputi Member)
U. grandis— Mor 1 1 1 Die odo (Morija), 3 indivs
Dicroidium open woodland (floodplain); Cycle 2c (Indwe Member)
Comparisons beyond Gondwana Triassic
Laurasian Triassic
Umkomasia franconia was described by Kirchner & Muller
(1992) from the Rhaeto-Liassic of the Bayreuth area (Gross-
bellhofen and Untemschreez), Germany. We believe this fructifica-
tion should be placed in a genus separate from Umkomasia. The
German material was associated with the leaf genus Thinnfeldia and
is clearly linked to that genus by the similar cuticle structure.
Other ages
An Umkomasia sp. has been recorded by Lejal-Nicol (1975)
from the Jurassic of Libya. Fragmentary pinnae from the same
locality were described as Dicroidium, Pachypteris and Thinnfeldia.
There are no generic characters available to confirm the presence of
Dicroidium leaves and we doubt the assignment of the ovulate struc- o
ture to Umkomasia. m
<
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
UMKOMASIALES
Umkomasia
244
d^TRELITZIA 15 (2003)
Molteno Umkomasia species, a comparative study
Umkomasia macleanii — llmk 111
Strobilus: small (ca 40 X 35 mm).
Megasporophyli. erect, bearing 2 or 3 pairs of pedicellate cupules; bracte-
oles usually in 2 pairs, at base and midway along peduncle.
Cupule: small, circular to ovate, unlobed.
Seed: circular, wing not evident.
U. bracteolata — Hla 213
Strobilus: small to medium (ca 55 X 30 mm).
Megasporophyli: erect, bearing a single pair of pedicellate cupules; bracte-
ole single, abaxial at base of peduncle.
Cupule: medium, circular to ovate, 2-lobed.
Seed: circular to ovate, wing narrow (incipient).
U. quadripartita — Mat 111
Strobilus: medium to large (ca 150 X 80 mm).
Megasporophyli: reflexed, bearing 3 or 4 pairs of pedicellate, partly decus-
sate cupules; bracteoles usually in 2 pairs, at base and midway along
peduncle.
Cupule: small to medium, relatively circular to ovate, 2-lobed (mainly).
Seed: circular to ovate, wing narrow to medium.
U. decussata— Aas 411
Strobilus: medium to large (to 7120 X 60 mm).
Megasporophyli: semi-erect, bearing up to 7 strongly decussate pairs of
sessile cupules; bracteole single, abaxial at base of peduncle.
Cupule: small, roundly triangular, 2-lobed (mainly).
Seed: circular, wing not evident.
U. monopartita— Mat 111
Strobilus: medium to large (>100 X 38 mm).
Megasporophyli: reflexed, bearing a single pair of pedicellate cupules;
bracteoles variable, usually single at abaxial base and paired midway
along peduncle.
Cupule: medium, roundly triangular, 4-lobed.
Seed: circular to ovate, wing medium.
U. gracilliaxis — Lit 111
Strobilus: large (>100 X 50 mm), distinctly lax; axis gracile.
Megasporophyli: erect to reflexed, bearing a single pair of pedicellate
cupules; bracteole single, elongated, abaxial at base of peduncle.
Cupule: medium to large, roundly triangular, 4-lobed.
Seed: ovate, wing narrow to medium.
U. cupulata—Kon 222
Strobilus: large (>100 X 40 mm), axis of medium girth.
Megasporophyli: reflexed, bearing a single pair of pedicellate cupules;
bracteole single, abaxial at base of peduncle.
Cupule: large, roundly triangular, 4- or 5-lobed.
Seed: ovate, wing narrow.
U. grandis— Mor 111
Strobilus: large (far >1 10 X 64 mm), axis robust.
Megasporophyli: reflexed, bearing a single pair of sessile cupules; bracteole
single, abaxial at base of peduncle.
Cupule: large, ovate, 2-lobed.
Seed: unknown.
Umkomasia
UMKOMASIALES
d^TRELITZIA 15 (2003)
245
Tab. 52. Umkomasia, Molteno occurrence
Evidence for affiliation of organs
Literature
Since Thomas (1933) described and affiliated Umkomasia
and Pteruchus with Dicroidium, this affiliation has been widely
accepted (Crane 1988; Retallack & Dilcher 1988) and is fully
supported by our research (And. & And. 1983, 1989).
In the last decade, excellently preserved permineralised fos-
sils have been described from one locality in the Fremouw
Formation in the Beardmore Glacier region of Antarctica by the
Taylors and their colleagues. This material links Dicroidium
leaves and stems with Pteruchus and Umkomasia on the basis of
vascular structure (Perovich & Taylor 1989; Pigg 1990; Meyer-
Berthaud et al. 1993; Osborn & Taylor 1993; Yao et al. 1995). A
recent paper on Antarctic material (Axsmith et al. 2000) is dis-
cussed under ‘Exclusions & synonyms’ above.
Mutual occurrence
In the Molteno, Umkomasia occurs at 22 TCs (Tab. 52) where,
in most cases, Dicroidium is a dominant element of the flora. The
remaining 53 TCs (Tab. 52) that have not yet yielded Um-
komasia are mostly under-collected (Tab. 1, man-hours collect-
ing). Umkomasia and Pteruchus have both been collected at 22
TCs. (A discussion on this co-occurrence is provided under
Pteruchus. p. 253.) At twoTCs (Cyp 111, Qua 111), Umkomasia
is present but Pteruchus has not been found. There is also the
puzzling affiliation of Fanerotheca (p. 274) and its apparent
association with Dicroidium at Bir 1 1 1 and Wal 111.
Cuticular correspondence
This was first noted by Thomas (1933) and more recently by
Townrow (1962) for Umkomasia, Pteruchus and Dicroidium.
Further details are given under ‘Cuticles’ (p. 240).
Species level affiliations
The question of which Umkomasia species affiliates with
which Dicroidium species is addressed only peripherally here,
and needs further resolution. We have made a preliminary corre-
lation between the Umkomasia species occurring at the 22 TCs
(Tab. 52) and the associated Dicroidium species (unpublished
foliage-species/TC matrix table; see also And. & And. 1983,
Tabs 6. 7; Figs 3-5, including ‘pictograph series’). At some six
TCs there is one dominant Dicroidium species (palaeodeme) and
only one Umkomasia species, but no consistent pairing between
the respective species emerges.
U. macleanii, occurring in eight TCs (and readily the most
frequent ovulate species), reveals no clear tie with a particular
Dicroidium species. It is most abundant at Umk 111 Die 2spp
(175 indivs) and could be linked to the Dicroidium palaeodeme
encompassing D. orbiculoides 25% and D. crassinervis 13%
(referred to as D. odontopteroides subsp. orbiculoides and D.
crassinervis forma obtusifolium and forma crassinervis respec-
tively in And. & And. 1983; see also Tabs 16, 17, this volume).
In spite of holding by far the largest (extensive-intensive)
collection of Umkomasia and Dicroidium from any formation
globally, we still do not have sufficient material to discern the
patterns of affiliation at a species level.
UMKOMASIALES
Umkomasia
246
C/TRELITZIA 15 (2003)
Umkomasia macleanii H.H.Thomas 1933
Holotype
Specimen: V. 23360 (U 11) in Nat. Hist. Mus., London.
Thomas (1933), fig. 1, pi. 23(56); refigured here, pi. 82(8).
Assemblage: Umk 111 Die 2spp, Umkomaas Valley.
Preservation: a virtually complete strobilus; compression in thinly lami-
nated, carbonaceous (good cuticle), moderately baked, dark grey shale
with good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 175 indivs (25 intact, 50 partial, 100 isolated), pi. 82(1-7,
10-16). (This does not include the Thomas (1933) collection of some
30 specimens in the Nat. Hist. Mus., London.)
Sister palaeodemes— 7 (best 3 listed)
Maz 211 Hei/Dic: 29 indivs (1 intact, 3 partial, 28 isolated), pi. 86(8-15).
Hla 213 Die elo; 28 indivs (3 intact, 7 partial, 18 isolated).
Lit 111 Dic/Hei: 9 indivs (all isolated).
Specific diagnosis
An Umkomasia species with small compact strobili, bearing erect
megasporophylls with 2 or 3 pairs of pedicellate cupules.
Specific characters
Strobilus: small ( ca 40 X 35 mm).
Megasporophyll: erect, bearing 2 or 3 pairs of pedicellate cupules; bracte-
oles usually in 2 pairs, at base and midway along peduncle.
Cupule: small, circular to ovate, unlobed.
Seed: circular, wing not evident.
Eponymy
macleanii — in honour of Tom Maclean, farmer in the area at the time of
Thomas’s trip to South Africa.
Comment & comparison
U. macleanii is the most frequently occurring (eight TCs) and abundant
of the eight Molteno Umkomasia species recognised here. The Umk 111
reference palaeodeme, with 175 individuals (including 25 more or less
intact strobili), is by far the most fully represented Umkomasia palaeodeme
from the formation. While Thomas (1933) recognised three genera with 14
species in his collection from the Umk 1 1 1 site, our interpretation of the
material is that it all falls within this single species. The 22 remaining indi-
viduals in our collection from Umk 111 are identified as U. monopartita
(Tab. 5 1 ) and have far larger fleshy appearing cupules and constitute a quite
distinct palaeodeme.
Umkomasia bracteolata j.M.And. & H.M.And , sp. nov.
Holotype
Specimen: PRE/F/8903a,b; pi. 83(1^1).
Assemblage: Hla 213 Die 3spp, Hlatimbe Valley.
Preservation: virtually complete strobilus, with counterpart; compression
in thinly laminated, carbonaceous (poor cuticle), medium dark grey
shale with good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 3 indivs (1 intact, 2 partial), pi. 83(1-8).
Sister palaeodemes— nil.
Specific diagnosis
An Umkomasia species with small to medium compact strobili, bear-
ing erect megasporophylls with a single pair of pedicellate cupules.
Specific characters
Strobilus: small to medium (ca 55 x 30 mm).
Megasporophyll: erect, bearing a single pair of pedicellate cupules; bracteole
single, abaxial at base of peduncle.
Cupule: medium, circular to ovate, 2-lobed.
Seed: circular to ovate, wing narrow (incipient).
U. bracteolata
Etymology
bracteolata— with reference to the bracteoles that are particularly well pre-
served.
Comment & comparison
This species, represented only by the very limited reference
palaeodeme of three individuals (including one intact strobilus), is the least
securely established of the eight Molteno Umkomasia species. It is disting-
uished from U. monopartita principally by size and the two-lobed rather
than four-lobed cupules.
Umkomasia
UMKOMASIALES
d?TRELITZIA 15 (2003)
247
Umkomasia quadripartita J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/9 1 57a,b; pis 84( 1,5), 85( 1 ).
Assemblage: Mat 1 1 1 Die dub, Matatiele.
Preservation: a virtually complete strobilus, part and counterpart; impres-
sion in thickly laminated, olive-grey shale with moderate cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 30 indivs (2 intact, 1 6 partial, 12 isolated), pis 84(1-5), 85( 1-6);
seeds, pi. 85(7-12), possibly (13-16).
Sister palaeodemes— 2 only (both listed)
Kra 111 Die odo: 13 indivs (1 intact, 3 partial, 9 isolated).
Nuw 1 1 1 Die zub: 11 indivs (1 intact, 4 partial, 6 isolated).
Specific diagnosis
An Umkomasia species with medium to large, relatively lax strobili,
bearing reflexed megasporophylls with 3 or 4 pairs of pedicellate cupules.
Specific characters
Strobilus: medium to large (ca 150 X 80 mm).
Megasporophyll: reflexed, bearing 3 or 4 pairs of pedicellate, partly decus-
sate cupules; bracteoles usually in 2 pairs, at base and midway along
peduncle.
Cupule: small to medium, relatively circular to ovate, 2-Iobed (mainly).
Seed: circular to ovate, wing narrow to medium.
Etymology
quadripartita— with reference to the four pairs of cupules per megasporophyll.
Comment & comparison
At Mat 111, the collection of 75 Umkomasia individuals (with eight
intact and 33 partial strobili) can be fairly readily separated into two
palaeodemes. These have been selected as reference for U. quadripartita
and U. monopartita respectively. With its multi-ovulate megasporophylls,
U. quadripartita compares most closely with U. macleanii but is distin-
guished in the greater number of cupule pairs and the far larger overall size.
Umkomasia decussata J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/20827a,b; pi. 86(1, 3).
Assemblage: Aas411 Dic/Sph, Aasvoelberg.
Preservation: a virtually complete strobilus, part and counterpart; impres-
sion in thinly laminated, strongly baked, yellowish grey shale with
very good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 7 indivs (1 intact, 3 partial, 3 isolated), pi. 86(1-7).
Sister palaeodemes— nil.
Specific diagnosis
An Umkomasia species with medium to large, relatively lax strobili,
bearing semi-erect megasporophylls with up to 7 strongly decussate pairs
of sessile cupules.
Specific characters
Strobilus: medium to large (up to ?120 X 60 mm).
Megasporophyll: semi-erect, bearing to 7 strongly decussate pairs of ses-
sile cupules; bracteole single, abaxial at base of peduncle.
Cupule: small, roundly triangular, 2-lobed (mainly).
Seed: circular, wing not evident.
Etymology
decussata— with reference to the opposite-decussate arrangement of the
cupules.
Comment & comparison
This extremely rare taxon, known only from Aas 41 1, is the most dis-
tinctive of the Molteno Umkomasia species in bearing sessile cupules in
strongly decussate pairs. This is seen clearly in the almost 3D preservation.
N I
I p™a
/ I Holotype
PRE/F/20827a,b
pi 86(1,3)
Holotype
UMKOMASIALES
Umkomasia
248
d^TRELITZIA 15 (2003)
Umkomasia monopartita J.M.And. & H.M.And., sp. nov.
Holotype
Specimen-. PRE/F/9175; pi. 87(1, 8, 9. 10).
Assemblage'. Mat 111 Die dub, Matatiele.
Presen’arion: incomplete strobilus, without counterpart; impression in thickly
laminated, olive-grey shale with moderate cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens'. 45 indivs (6 intact, 17 partial, 22 isolated), pi. 87(1-13).
Sister palaeodemes— 4 (best 1 listed)
Umk 111 Die 2spp: 22 indivs (5 partial, 17 isolated).
Specific diagnosis
An Umkomasia species with medium to large, relatively lax strobili,
bearing reflexed megasporophylls with a single pair of pedicellate cupules.
Specific characters
Strobilus : medium to large (>100 X 38 mm).
Megasporophyll: reflexed, bearing a single pair of pedicellate cupules;
bracteoles variable, usually single at abaxial base and paired midway
along peduncle.
Cupule: medium, roundly triangular, 4-lobed.
Seed: circular to ovate, wing medium.
Etymology
monopartita— With reference to the single pair of cupules per megasporophyll.
Comment & comparison
This well represented species shares a number of features with U.
gracilliaxis and differs mainly in its more compact strobilus with shorter
pedicels.
Umkomasia gracilliaxis J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: BP/2/2110; pi. 88(1, 3).
Assemblage: Lit 111 Dic/Hei, Little Switzerland.
Preservation: incomplete strobilus, without counterpart; compression in
thinly laminated, carbonaceous (good cuticle), dark grey shale with
moderate cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 42 indivs (1 intact, 10 partial, 31 isolated), pi. 88(1-13).
Sister palaeodemes— 3 (best 1 listed)
Maz 211 Hei/Dic: 17 indivs (2 intact, 6 partial, 9 isolated).
Specific diagnosis
An Umkomasia species with large lax strobili, bearing erect to reflexed
megasporophylls with a single pair of pedicellate cupules.
Specific characters
Strobilus: large (>100 X 50 mm), distinctly lax; axis gracile.
Megasporophyll: erect to reflexed, bearing a single pair of pedicellate
cupules; bracteole single, elongated, abaxial at base of peduncle.
Cupule: medium to large, roundly triangular, 4-lobed.
Seed: ovate, wing narrow to medium.
Etymology
gracilliaxis— with reference to the gracile axis of the strobilus.
Comment & comparison
U. gracilliaxis has a particularly gracile and lax strobilus and it is main-
ly in this feature that it differs from all other Molteno Umkomasia species.
It is closest in overall morphology to U. monopartita and U. cupulata.
Umkomasia
UMKOMASIALES
c/tRELITZIA 15 (2003)
249
Umkomasia cupulata J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: BP/2/41 22a. b; pi. 89(1, 2).
Assemblage: Kon 222 Die odo, Konings Kroon.
Preservation: fragmentary strobilus, part and counterpart, 3D mould and
cast in thinly laminated, medium grey cherty shale with poor cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 20 indivs (5 partial, 15 isolated), pi. 89(1-5).
Sister palaeodemes-
Specific diagnosis
An Umkomasia species with large relatively lax strobili, bearing
reflexed megasporophylls with a single pair of pedicellate cupules.
Specific characters
Strobilus: large (>100 X 40 mm), axis of medium girth.
Megasporophyll: reflexed, bearing a single pair of pedicellate cupules;
bracteole single, abaxial at base of peduncle.
Cupule: large, roundly triangular, 4- or 5-lobed.
Seed: ovate, wing narrow.
Etymology
cupulata— in recognition of the clear preservation of the cupules.
Comment & comparison
U. cupulata , known only from Kon 222, comes closest to U. grandis
in size but the cupules are pedicellate (not sessile), roundly triangular (not
oval) and split into four or five lobes (not two).
UMKOMASIALES'
Umkomasia grandis J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/3397; pi. 89(6, 7).
Assemblage: Mor 111 Die odo, Morija.
Preservation: intact strobilus, no counterpart, impression in thickly lami-
nated, light olive-grey shale with poor cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 3 indivs (1 intact, 2 isolated), pi. 89(6-8), p. 245, tf. 1.
Specific diagnosis
An Umkomasia species with large, robust, relatively lax strobili, bear-
ing reflexed megasporophylls with a single pair of sessile cupules.
Specific characters
Strobilus: large (far >1 10 X 64 mm), axis robust.
Megasporophyll: reflexed, bearing a single pair of sessile cupules; bracteole
single, abaxial at base of peduncle.
Cupule: large, ovate, 2-lobed.
Seed: unknown.
Etymology
grandis— with reference to the large size of the strobilus.
Comment & comparison
U. grandis, in life, would have been by far the largest of the Molteno
Umkomasia species. It may easily have reached an impressive 0.3 m in
length. The species differs from U. cupulata in its far greater overall size
and in the megasporophylls bearing single pairs of sessile cupules which
split into two lobes (not four or five).
Umkomasia
250
<§?'. TRELITZIA 15 (2003)
Pteruchus H.H.Thomas 1933
Type species
Pteruchus africanus H.H.Thomas 1933.
Umkomaas Valley, Karoo Basin, S. Africa; Camian, Triassic.
Generic diagnosis emended
A ginkgoopsid male strobilus of lax paniculate form, with simple to
once-forked microsporophylls bearing terminal laminate heads with
numerous abaxial microsporangia.
Generic characters
Strobilus: simple, lax, paniculate, radially symmetrical, small to large (30-200
mm long); axis robust, markedly tapering, erect to arching; microsporo-
phylls fairly numerous, arrangement irregularly helical.
Microsporophyll: simple to once forked near base, planar; peduncle gracile;
fertile heads single or paired, pedicellate, oval to linear oblong (5 — 40
mm long); laminae bilaterally symmetrical, margins undulate to strong-
ly lobed; microsporangia numerous, pendent, densely packed to cover
full abaxial surface of lamina.
Microsporangium: linear-elliptic (2-4 mm); ornamentation fine, linear, fork-
ing and converging.
Pollen: disaccate.
Etymology
Pteruchus— pterona (Gr). wing; Thomas (1933) described the microsporophyll
as ‘a wing-like thing’.
Global range: several spp., Gondwana, Tr. (SCY-CRN).
First: Pteruchus feistmanteli (Pterorrachis barreal) Retallack ( 1973).
Mt. Piddington, Banks Wall Fm., Blue Mts, Australia.
Last: Pteruchus africanus ( Baiera tenuifolia ) Johnston (1888).
Lord’s Hill. Brady Fm., Hobart, Tasmania.
Gondwana Triassic occurrence (see Pteruchus hypodigm. Tab. 53)
SAm— N. Argentina & Patagonia, 5 TCs (12 indivs).
SAf— Karoo Basin. 22 TCs (431 indivs).
Ind— S. Rewa/Tiki, 1 TC (1 indiv.).
Ant— E. & Trans- Antarctica, 3 TCs (11 indivs).
Aus— Australia widespread, 13 TCs (33 indivs).
New Zealand, 2 TCs (2 indivs).
Molteno occurrence (see Tab. 54)
Frequency (F): 22 TCs (of 100 sampled in the Molteno).
Diversity (D): 3 species.
Abundance (A): 431 indivs total; rare to very rare in top 8 TCs.
Kra 111 Die odo: 30 indivs in 20 man-hrs cleaving (15 per 1 man-day) rare
Mat 1 1 1 Die zub:
84
” 65
(14 ” 1
” ) ”
Nuw 111 Die zub:
15
” 21
”
( 7 ” 1
” ) ”
Kon 222 Die odo:
13
” 40
( 3 ” 1
” ) ”
Hla 213 Die elo:
20
” 60
( 3 ” 1
” ) ”
Maz 211 Hei/Dic:
27
” 85
( 3 ” 1
” ) ”
Umk 111 Die 2spp:
138
” 400
”
( 3 ” 1
” ) ”
Lit 111 Dic/Hei:
36
” 550
( 1 ” 2
” ) v. rare
As for Umkomasia, we include above only those eight TCs yielding
the greatest number of Pteruchus specimens. The pattern of occurrence of
the reproductive affiliates of Dicroidium appears remarkably similar.
Affiliated organs
Female strobilus: Umkomasia— Grade 4 (Mut. occ.. Cut. corn).
Foliage: Dicroidium— Grade 4 (Mut. occ.. Cut. corn).
Classification & comparison
Suprageneric classification (Umkomasiaceae/Umkomasiales)
A group of six Gondwana Triassic male genera are included here in the
Ginkgoopsida: Antevsia (Peltaspermales), Switzianthus (Matatiales), Eosteria
(Ginkgoales), Stachyopitys (Hamshawviales), Pteruchus (Umkomasiales)
and Kannaskoppianthus (Petriellales). Based on their female affiliates,
which all differ significantly in morphological organisation, each is placed
in a separate family and order.
Intergeneric comparison (Gondwana Triassic)
Of the group of six genera noted above, Antevsia , Stachyopitys and
Pteruchus are most alike in that each bears clusters of similarly elongated
microsporangia. Poorly preserved specimens are easily confused.
However, Pteruchus is differentiated by the clear laminate head bearing
numerous abaxial microsporangia; Stachyopitys has numerous micro-
sporangia radiating out from a central receptacle; and Antevsia has sessile
fascicles of microsporangia. The remaining three genera ( Switzianthus ,
Eosteria, Kannaskoppianthus) each display an entirely different organisa-
tion.
Reconstructions
Strobilus
Certain reconstructions of the Pteruchus strobilus (Crane 1985, p. 755;
Retallack & Dilcher 1988, p. 1042) show a dorsiventral structure with the
microsporophylls arranged in a single plane. From a study of perminer-
alised specimens of Pteruchus from Antarctica, Yao et al. (1995) provide a
reconstruction that shows helically borne microsporophylls. With some
reservation, we believe that the Molteno Pteruchus material also has an
irregularly helical arrangement as illustrated opposite.
Microsporophyll
The size of the microsporophyll and whether it is single or paired vary
according to species. The marginal lobing of the lamina possibly occurs in
all species, but is often obscured by the numerous microsporangia. The lob-
ing was noted by Thomas (1933, p. 234) and Townrow (1962, p. 294, tf. 2E)
and is clearly seen in many specimens in our collection, e.g. pi. 94(8, 9).
As shown in our reconstructions of P. matatimajor opposite (tfs. 1, 2),
a subtending bracteole occurs at the base of each pedicel— clearly seen also
in pis 92(9, 11), 93(1, 6). Bracteoles probably occurred on most specimens,
but may have been lost before or during preservation. As noted by
Townrow (1962, p. 292), who termed them 'vegetative pinnules’, they can
be quite inconspicuous.
Gondwana Triassic occurrence (elaborated)
Pteruchus , after Stachyopitys, is the second most frequent (22 TCs)
and abundant (425 indivs) pollen-bearing genus in the Molteno (Tabs 6b,
9b and 1 1 ) and the most widely recorded through the Gondwana Triassic.
Though Pteruchus is now known from all five Gondwana continents (Tab.
53), its true preserved spread and abundance may be strongly under-report-
ed. The most extensive collections come from South Africa (Tab. 54) fol-
lowed by eastern Australia.
The most widespread species are P. barrealensis and P. feistmantelii,
followed by P. africanus. However, most published specimens are insuffi-
ciently complete to identify to species level (see Tab. 53, Pteruchus hypo-
digm).
Pteruchus
UMKOMASIALES
T RELITZIA 15 (2003)
251
UMKOMASIALES
Pteruchus
252
dA'RELITZIA 15 (2003)
Tab. 53
PTERUCHUS HYPODIGM, Gondwana Triassic occurrence
Species
Intact-
ness
Molteno
Other
P. africanus
P. matatimajor
P. helvetigracilis
I'll
^2 1 r”
® 1 3
c' </>' d
a; ; a: ; o:
Intact strobilus
Fragm. strobilus
Isolated heads
AUTHOR
SUBREGION
FORMATION
LOCALITY
NAME
Indivs ILLUSTRATION
SOUTH AMERICA
1876 (Geinitz
Marayes NA3
28 Q. de la Mina
Marayes (Huerta Mts)
Sphenolepis rhaetica
2 ! pi 2(23-24)
j |
- 1 . '?2
jL 1 ; 1
1942 Frenauellli
Barreal NA2
22! Barreal
Punto II ?
Pterrorrachis barrealensls
1 ; p 2
.1 _ ! .
1 ' - ' -
1 1 - -
1944a ” (specimen repeated from Frenquelli 1942)
Zubeha zub. (Pteror. barreal.)
* < pi 11(1-5)*
-1- ! .
* | J .
1956 Pinto
S. Maria PA1
28 S. Maria
Passo das Tropas
Pteruchus
1 I pi 2(3)
I |
-1 - !?i
-Ii!-
1963* Bonetti
Barreal NA2
23! Cortaderita
Punto 32
" dubius
1 ! pi 26
.
rJ.il-
1967 Jain & Del.
Cacheuta NA4
24! Potrerillos
Minas de Petroleo
” rhaetica
1 ! pi 90(15)
1 1
. 1 . 1 _
1
" Bort. & Barb, (specimen repeated from Pinto 1956)
Pteruchus
* | Pi 2(3)*
_ 1 _ l *
1980 Petriella (specimen repeated from Frenguelli 1942)
" dubius var. barreal.
* • pi 1(1)*
* 1 1
* !
” (specimen repeated from Bonetti 1963b)
*Tpiim*
*1' 1
. \ * j .
n | ..
? ) ?
? 1 ?
?
3 pi 1(3-5) .
3 1 - 1 -
-! -! 3
? ; ?
?
?
" simmondsi
3 ! pi 2(1-3)
- ’ - !?3
- ! - ! 3
SOUTH AFRICA
1933 Thomas
Underberg 1 Ka4
24 Molteno
Umkomaas
Pteruchus africanus
2 ! p| 24(71-72), tf 34-35
2! - ! -
1 I
ill!-
" J -
" papillatus
1 1 tf 36-37
1! -! -
1 ! -! -
- «
” p eltatus
1 ! tf 38-39
1 ! - ! -
- ' . 1 .
" j ”
" hoeai
1 1 pi 24(75), tf 40-41
1; -; -
. ! .! .
1 1 - 1 -
" ;
" stormberpensis
1 1 tf 43
V -' -
1 1 - 1 -
” ,
" dubius
1 ! tf 44-45
ii -1 -
. | .
-I -Ii
,, |
” !
" minor
1 ! pi 24(76), tf 46
1! -! -
-: i| -
" | "
” 1
" sp. Type X
1 ! tf 47-48
_ 1 . » .
- 1 1 ! -
1962 Townrow
" africanus
6 ; pi 24(4), 25(1,2),
6; - ; -
.I.i.
114
- 1 26(2,4-11), tf 1(A-D,F)
.1.1.
7TT1T
| |
- ! 2(D-G), 3(B-C)
" ) ”
" dubius
2 pi 24(5,6), 25(3),
2! -I -
1 1 - ; 1
1
- ; tf 1(E), 4(B,C)
i 1
. | . 1 .
” |
” | "
j
” simmondsi
1 ; pi 24(1,3), tf 2(A-C)
1; - : -
.i.i.
1 ! -' -
” | "
"
- 1 4(A,D-H)
.1.1-
. 1 . 1 .
1 J
1978-1999: And.& And. Molteno literature not included in this table
! i
INDIA
1962b Lele
S. Rewa/Tiki PL1
12
Kamtadand (loc. 9)
Marattiopsis sp. A
1 : pi 2(16.17) tf 3
- - -
?i» - j -
ANTARCTICA
1989 Taylor ef al.
Queen Alexander 1 TA4
19! Fremouw
Fremouw Peak
Dicroidium pollen organs
1 ! tf 9
| |
-! -i?i
-I -1 1
1993 Osb. & Taylor
” j »
-■ ;
Corystosperm pollen sacs
i_L.pi 1
- L-
-1-1 1
1 995 ! Yao ef al.
Pteruchus fremouwensis
6 1 f (1-?6)
.' . 1 .
- < - ;?i
- : 1 ■ 4
1995 Cantrill ef al.
Pr. Charles Mts 1 EA1
? 1 Flagstone Beach
PCM 19
" dubius
2 1 f 5 (a,c)
i.i.
- 1 - 1?2
-! -I 2
2000 Axsmith ef al.
Shackleton Gl. TA6
? i ?Fremouw
Alfie’s Elbow
Pteruchus
2 ! f 19-29
?2 - ' -
-■ ! - ! .
2! - 1 -
NEW ZEALAND
1980a Retallack
Mt Potts ! N23
21 Tank Gully CM
Tank Gully
" johnstonii
1 | f 5A
?l! - ! -
-! 1' -
1983 : "
Benmore Dam , N24
21 : Bl. Jack Conql.
Benmore Dam
" dubius
1 ' f 6A
-' - 1 -
-! - '?i
- ! - 1
AUSTRALIA
1888 Johnston
Hobart Ta5
25! Brady equiv.
Lord's Hill (Hobart)
Baiera tenuifolia
1 ! pi 27(2D,E)
?il - ! -
1 i - ; -
1890b Feistmantel (s
1898 Shirley
pecimen repeated from Johnston 1888)
Trichopitys johnstonii
* ; pi 10(5)*
*; -1 .
Ipswich/Esk ICM5 24 Blackstone Stage
Denmark Hill
Stachyopitys annularioides
1 1 pi 18(1)
.'.I.
. \ . i??i
1 ; - 1 -
1925b Walkom (specimen repeated from Johnston 1888)
* ! f 15-16*
*! . ! .
1947 Jones & de J.
Ipswich/Esk i CM5
23i Tingalba
Campbell's Quarry
Pteruchus annulariodes
1 ! tf 38
- 1 - 1 -
-! - !?i
1 - ! 1
" [ ”
19 1 Blackstone Stage
Parish Goodna
” africanus
1 ! tf 51
?i; - ; -
" | ”
24 Blackstone Stage
Denmark Hill
Strobilus
1 ! tf 53
- 1 - ;?i
-I 1
1948 Jones
Bowen Basin ! ?
? ; ?
Walhalla, Cracow
Strobilus
1 ' pi 1(15)
. 1 .
- ! -'?i
-j - j 1
1962 Townrow (specimen repeated from Johnston 1888)
Pteruchus johnstonii
* I tf A-C*
*! . ! .
. | -i .
1965 Hill ef al.
Ipswich/Esk | CM5
24 Blackstone Stage
Denmark Hill
” dubius
1 ! pi T6(3)
.! . ! .
?i! - -
1 i - ! -
24 Blackstone Stage
Denmark Hill
” johnstonii
1 ! pi T6(4)
-!-:i
1 ! - ; -
1973* Retallack
Blue Mts. * Sy2
8 Banks Wall
Mt. Piddington
' Pterorrachis' barreal.
2 ! pi 16(3), f 17B,
- 1 2; -
-! 2' -
Sydney i Sv3?
? I ?
?
7 ' pi 16(1.2.4-7). f 17JA.D)
- : 7 1 -
- ! 4 1 3
1974 Pledge
?Springfield ] SA?
22| ?
?
? Pteruchus
1 pi 7(5) .
.i.i.
- l - 1?1
- 1 - 1
1977 Retail, ef al
Nymboida j CM7
19| Cloughers Cr.
UNEL 1564
Pteruchus johnstonii
1 ! f 8 (C)
.! . ! .
- ! - ,?i
-1 1! -
(Kangaroo Creek)
" dubius
1 ! f 8 (D)
. 1 . ! .
- ; -!?i
-1 1
1979 Holmes & Ash
Lome B NE2
9 1 Camden H. Cl.
Camden Head (1583)
" barreal. var. feistm.
3 ! f 6 (9-11)
- 1 3; -
-! -'3
1979* Rowett
Leigh Creek SA1
- ! ?
Telford B Lobe B
_
74' pi 24(1-4)
- 1 - '?4
- ! 1 1 3
1980b Retallack (specimen repeated from Retallack 1973)
'Pterorrachis' barreal.
* ! f 21.9(D)*
.i.i.
1981a
Sydney I Sy3
? 1 Newport
?
1 ! f 10-4(C)
-! i.i -
- ' 1 ! -
" \ "
? ’
?
Umkomasia sp.
1 ! f 10-4(D)
-; - !?i
- . i_Lr
1982 ‘Holmes
Dubbo (Sy1
18 (unnamed)
Benolonq (Uqotherv)
Pteruchus johnstonii
2 ; f 7 (E-F)
2! - ! -
2! - 1 -
” : Playf. ef al.
Canarvon i Bo5
1 9 1 Moolayember
Spring Creek
" simmondsii
1 1 pi 2(5-6)
- 1 -
- ! - >?i
J 1! -
1 984 i Taylor ef al.
Ipswich/Esk ;CM5
? ! ?
Dinmore
” dubius
1 ! pi 1(1-2)
-I.I.
?1 - ; -
ifl -
1986 White (specimen repeated from Holmes & Ash 1979)
’’ barrealensis
* 1 phon p 148 (220)*
-j -|-
* unpublished
* specimens repeated — number of individuals therefore not given
Pteruchus
UMKOMASIALES
TRELITZIA 15 (2003)
253
assemblages
(taphocoenosis)
Ken 111 Die era
Nav 111 Die odo
Cal 111 Dic/Sph
Bir 211 Sph 2spp
" 311 Hei/Sph
Bir 111 ”
Gre 121 Hei elo
111 Equ sp.
Boe 111 Lep sto
” " Dic/Hei
” 112 Die cor
Cyp 111 Die era
Mol 111 Sph pon
Kan 112 Hei elo
Tel 111 ” ’’
Kom inSph/Dic
Vin 111 Die odo
Ela 111 ” "
Kra 311 ” ”
” 221 Beetles
" 111 Die odo
Lut 111 Hei/Dic
” 411 " ’’
" 311 Hei elo
" 221 Equ sp.
Tin 121 Sph 2spp
Wal 111 Die odo
Kon 223 " ’’
” 222 ” ”
" 211 Hei elo
’’ 111 Die odo
" Hei elo
Pen 321 Dic/Ris
211 Dic/Equ
" 221 " "
" 511 Equ sp.
” 421 Die odo
" 431 Dic/Equ
” 311 Hei elo
” 411 " ”
Kle 111 Hei/Dic
Kap 111 Dic/Ris
Ela 112 Equ sp.
Dic/Hei
Nuw 111 Die zub
” 211 Pic 2spp
Win 111 Hei elo
Mor 111 Die zub
” Die odo
Qua 111 Die odo
Mak 111 " ’’
Maz 111 Die era
’’ 211 Hei/Dic
Moo 111 Die zub
Hla 111 Equ sp.
" 211 Die 3spp
" 212 " ”
” 213 Die elo
Umk 111 Die 2spp
Cha 111 Die odo
” 211 Die dub
Inj 111 Die odo
" 211 Die dub
San 111 Die era
Mng 111 Die 2spp
Qac 111 Hei/Dic
Mat 111 Die dub
Gol 111 " ”
Lit 111 Dic/Hei
Aas 111 Hei elo
” 311 ’’ "
”411 c/Sph
” 511 Die elo
Ask 111 Equ sp.
Bam 111 Pic dub
Total TCs
Total indivs
Genera
c S 3 «
fc E -C ■£
•3' o 3 2
■a 2 *
5 E a §
b => 0- u-
S 1 9 1 cf 1 9
_L ‘
2: 1 2
- - 60
75. 22. 22 27
%! 503425 .247
Species Intactness
Q. r o. J£
10
-! i
5 5
-' 11' 19
TIT!
ii ' -i
15
1 i - ' AT
27
17 i -;
1138 -'
“9 '15! 3
i; i; -
1. T 2
3 11 6
34 64 40
17 l_4i 1 i
327! 91 T 3l
15 34 35
t; -!"-
3' 19 14
10 15 21
70 !l 89 466
-I L
Tab. 54. Pteruchus, Molteno occurrence
Comparisons beyond Gondwana Triassic
Laurasia Triassic
Besides Stachyopitys and Antevsia , no other genera with a close-
ly similar structure to Pteruchus are known to us.
Pteruchus septentrionalis , together with the doubtful Umkomasia
franconia (p. 243), has been described by Kirchner & Muller (1992)
from the Bayreuth area (Grossbellhofen and Unternschreez),
Germany. The Pteruchus specimen illustrated by them on pi. 3(4) is
possibly close to Pteruchus, while that on pi. 3(2) is something very
different. We do not accept these as Pteruchus and suggest that they
belong to a new genus affiliated to Thinnfeldia to which they are
clearly linked by the similar cuticle structure.
Other ages
Certain male strobili from the Carboniferous, e.g. Crossotheca
and Feracotheca (order Lagenostomales), bear numerous abaxial
microsporangia and in this way show similarities to Pteruchus (see
Taylor & Taylor 1993).
Evidence for affiliation of organs
The female /male balance in occurrence (Molteno Fm.)
The pattern of occurrence of the female and male strobili affiliating
with Dicroidium is quite unique. It is the only case, for the 20 gymnosperm
ovulate genera recognised within the Molteno, where the pattern of distribu-
tion (presence and abundance) of the affiliated organs appears closely par-
allel. This parallelism of occurrence shows very clearly in Tabs 52, 54. Um-
komasia and Pteruchus each occur in 22 TCs, and appear together in similar
abundance in most cases (in 20 TCs).
In marked contrast, a strong imbalance in preservation between male
and female organs is the normal pattern: Dordrechtites and Fraxinopsis,
for instance, are frequent and abundant, but for neither is there any evident
male counterpart; Telemachus and Peltaspermum are very much more fre-
quent and abundant than their male affiliates, Odyssianthus and Antevsia,
respectively; Rissikistrobus and Rissikianthus, female and male cones of
Rissikia, generally co-occur but either the one or the other is distinctly
more abundant; in Kannaskoppia, where the reverse to the general pattern
is observed, the male strobilus far outstrips the female in frequency; and
lastly, in a few cases, male cones occur without any apparent female coun-
terpart.
Why is the Umkomasia! Pteruchus pattern unique? What is there about
the Dicroidium plant that makes it different from all other Molteno gymno-
sperms in this regard? Why should it be the only case where the effect of
taphonomic filtering appears to be so strikingly similar for female and
male? We need, perhaps, to look at the other unique facts about Dicroidium.
It is by far the most prominent/successful gymnosperm genus in Gondwana
with regard to the combination of ubiquity, frequency, abundance and
diversity (And. & And. 1989; pp. 46, 68). It is the dominant genus in three
of the seven recognised Molteno habitats (Caimcross et al. 1995):
Dicroidium riparian forest (mature and immature) and the Dicroidium open
woodland of the floodplain. Is the explanation behind its success in some
way related to that behind its uniquely parallel female/male pattern of
occurrence?
Adaptive radiation (Molteno diversity)
The numerous species described by Thomas (1933) and Townrow
(1962) from Umk 1 1 1 (Umkomaas Valley) are all considered to be P. afri-
canus (see Pteruchus hypodigm. Tab. 53).
While 19 species of Dicroidium and eight species of Umkomasia— the
well-established foliage and female affiliates of Pteruchus respectively—
are recognised for the Molteno, only three species of Pteruchus can be
readily differentiated. The diagnostic characters of Pteruchus lie primarily
in the shape and size of the fertile heads, the nature of the lamina margin,
and whether or not the pedicels are forked.
The three species, each based on substantial palaeodemes from a dif-
ferent TC (all from the Indwe Member) and derived from Dicroidium ri-
parian forest habitats, are:
P. africanus— Umk 111 Die 2 spp (Umkomaas Valley), 138 indivs
Dicroidium riparian forest (mature); Cycle 2b (Indwe Member)
P. matatimajor—Msit 111 Die dub (Matatiele), 84 indivs
Dicroidium riparian forest (immature); Cycle 2b (Indwe Member)
P. helvetigracilis— Lit 111 Die Hei (Little Switzerland), 3 indivs
Dicroidium riparian forest (immature); Cycle 2a (Indwe Member)
UMKOMASIALES
Pteruchus
254
d^TRELITZIA 15 (2003)
Pteruchus africanus H.H.Thomas 1933
Holotype
Specimen : V23384 (U244), Nat. Hist. Mus., London.
Thomas (1933), fig. 34, pi. 24 (71); refigured here, p. 255, tf. 4.
Assemblage (TC): Umk 111 Die 2spp; Umkomaas Valley.
Preservation : an intact strobilus, compression in thinly laminated, car-
bonaceous (good cuticle), moderately baked, dark grey shale with
good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens'. 138 indivs (34 intact, 64 partial, 40 isolated), pi. 90(1-9).
Sister palaeodemes— 16 (best 2 listed)
Hla 213 Die elo: 17 indivs (2 intact, 9 partial, 6 isolated), pi. 91(1-6).
Kra 111 Die elo; 30 indivs (11 partial, 19 isolated).
Specific diagnosis
A Pteruchus species with small strobili, bearing unforked microsporo-
phylls with oval fertile heads and relatively robust microsporangia.
Specific characters
Strobilus: small ( ca 30^10 mm long).
Microsporophyll: peduncle unforked.
Fertile head: oval (ca 5-10 mm long).
Microsporangium: relatively robust.
Etymology
africanus— named by Thomas (1933) as coming from Africa.
Comment & comparison
Though there is some apparent morphological overlap between this
species and P. matatimajor at the smaller end of the latter’s range, the two
species are clearly distinct. The full collection of 138 Pteruchus specimens
from Umk 1 1 1 constitutes a single palaeodeme of small strobili with small,
unpaired, circular to oval microsporangial heads.
Pteruchus matatimajor j.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/9229a,b; pi. 92(5, 12).
Assemblage (TC): Mat 111 Die dub, Matatiele.
Preservation: central part of strobilus with tip and base missing, part and
counterpart; impression in thickly laminated, olive-grey shale with
moderate cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 84 indivs (15 intact, 34 partial, 35 isolated), pis 92-94.
Sister palaeodemes— 3 (all listed)
Hla 213 Die elo: 1 indiv. (1 intact), pi. 91(7-10).
Kon 222 Die odo: 3 indivs (3 partial).
Aas 41 1 Dic/Sph: 3 indivs (1 intact, 2 isolated).
Specific diagnosis
A Pteruchus species with medium to large strobili, bearing forked
microsporophylls with oblong to linear-oblong fertile heads and relatively
robust microsporangia borne on deeply lobed leafy laminae.
Specific characters
Strobilus: medium to large (ca 45->120 mm long).
Microsporophyll: peduncle normally once forked.
Fertile head: oblong to linear-oblong (5-25 mm long); leafy lamina with
deeply defined lobes.
Microsporangium: relatively robust.
Etymology
matatimajor— with reference to the large size of the specimens from the
Matatiele locality.
Comment & comparison
The large Mat 1 1 1 Pteruchus collection of 84 individuals covers a
wide range of variation in size, yet appears to constitute a single
palaeodeme characterised by the consistently forked microsporophylls and
linear-oblong fertile heads. Particularly distinctive are the laminae with
their coarse, clear lobing [pi. 94(8, 9)].
Pteruchus
UMKOMASIALES
g/tRELITZI A 15 (2003)
255
Pteruchus helvetigracilis j.M.And. & H.M.And., sp. nov.
Holotype
Specimen : BP/2/5793a,b; pi. 95(1-3, 5-7).
Assemblage (TC): Lit 111 Die Hei, Little Switzerland.
Preservation : an incomplete microsporophyll, part and counterpart; com-
pression in thinly laminated, carbonaceous (good cuticle), dark grey
shale with moderate cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 3 indivs (3 isolated), pi. 95(1-7).
Sister palaeodemes — nil.
Specific diagnosis
A Pteruchus species of unknown size, bearing microsporophylls with
linear fertile heads and relatively gracile microsporangia.
Specific characters
Strobilus: unknown.
Microsporophyll: peduncle unknown.
Fertile head: linear (>30 mm long), margins of leafy lamina uncertain.
Microsporangium: relatively gracile.
Etymology
helvetigracilis— helved (Lat.), Switzerland, referring to the type locality
Little Switzerland; gracilis (Lat.), slender, referring to the form of the
microsporangia.
Comment & comparison
With only three incomplete microsporangial heads from Lit 111, this is
by far the rarest and most infrequent of the Molteno Pteruchus species. It
could possibly be included with P. matatimajor , both seemingly affiliated
with Dicroidium zuberi, but does appear distinctly more gracile in all
respects.
Cuticles
Potential sample: Lit 111, 36 indivs; Umk 111, 138 indivs.
Macerated: none in this work; see Thomas (1933) and Townrow (1962).
Preservation grade: Grade 4-5.
Diagnostic characters: cells isodiametric to linear-oblong, end walls
square to oblique, walls gently curved to sinuous, nonpapillate to
papillate; trichome bases present; stoma orientation (?)random; sub-
sidiary cells brachyparacytic and actinocytic, nonlappetate; guard cells
narrowly elliptic.
Comment: —
Significance:
Classification— The cuticles of Pteruchus , as described and illustrated
by Thomas (1933) and Townrow (1962), are closely similar to those of
Umkotnasia (in Thomas 1933) and Dicroidium (in And. & And. 1983,
1989)— thus supporting classification in the order Umkomasiales.
Affiliations— As noted for Umkotnasia (p. 240), and for the same rea-
sons, the cuticle of Pteruchus clearly supports affiliation with Dicroidium
rather than the other Molteno ginkgoopsid foliage genera Lepidopteris,
Sphenobaiera, Ginkgoites and Dejerseya.
UMKOMASIALES
Pteruchus
256
d/TRELITZIA 15 (2003)
Dicroidium Gothan 1912
Type species
Dicroidium odontopteroides (Morris 1845) Gothan 1912.
Jerusalem Basin, Tasmania; Camian, Triassic.
Generic diagnosis
A Y-shaped ginkgoopsid leaf with a short distinct petiole and simple to
tripinnate lamina with lateral venation arching and forking.
Generic characters
Sterile shoot : with terminally attached fascicles of 2-5 or more leaves.
Leaf. Y-shaped, medium to large, ca 200-300 (up to 1000 mm) long;
rhachis dichotomising about a third from base; lamina entire to tripin-
nate; petiole short; pinnae contracted or rarely decurrent at base, highly
variable in shape; venation varying with pinna shape from taeniop-
teroid to odontopteroid with veins arching and forking once or twice.
Cuticle: see And. & And. (1989, p. 69); this vol., tfs 5a-c.
Etymology
Dicroidium— dichro, dikros (Gr.), forked; idium (Lat.), a suffix indicating
diminutiveness.
Global range: 21 spp., Gondwana, Tr. (SCY-NOR).
First: Dicroidium zuberi (Thinnfeidia odontopteroides) (Feistmantel, 1890);
Banks Wall Fm., Mt. Piddington, Blue Mts, Australia.
Last: Dicroidium odontopteroides ( Thinnfeidia lancifolia ) (Solms-Laubach
& Steinmann 1899); Quebr. La Temera, Copiapo, Chile.
Gondwana Triassic occurrence
Frequency (F): 45 degree squares (of the 84 across Gondwana).
Ubiquity (U): 5 continents (of 5 comprising Gondwana).
Diversity (D): 21 species.
Abundance (A): 90% (the norm in preferred Molteno habitats).
Longevity (L): 27 myrs (Smithian to late Norian).
Colonisation success: FUDAL rating 45/5/21/90/27 = 188.
Maximum success (Grade 5); Dicroidium was clearly the most prominent
genus in the Gondwana Triassic; it was ubiquitous, frequent, diverse,
abundant and long-lived.
Endemism: many of the Dicroidium species and infraspecific taxa colonised
widely across Gondwana.
Molteno occurrence
Frequency (F): 75 TCs (of 100 sampled in the Molteno).
Diversity (D): 21 species.
Abundance (A): monodominant (>70%) in 32 TCs; co-dominant (20-69%)
in 23 TCs; occasional to abundant (1-19%) in 12 TCs; rare to very rare
(<1%) in 8 TCs.
Habit: woody, probably from shrubs to large canopy trees.
Preferred habitat: the dominant genus in 3 of the 7 primary Molteno habi-
tats— Dicroidium riparian forest (types 1 and 2) and Dicroidium wood-
land.
Affiliated organs
Female strobilus: Umkomasia— Grade 4 (Mut. occ., Cut. corn).
Male strobilus: Pteruchus— Grade 4 (Mut. occ., Cut. com).
Classification & comparison
See And. & And. (1989, p. 68).
Adaptive radiation (diversity)
Dicroidium , a well-defined genus, is particularly diverse at the species
and infraspecific level. It is primarily in the shape of the pinnae and pin-
nules that the different taxa are recognised. Evidence suggests that the
Umkomasia! Dicroidium plant-genus actively colonised and diversified
through much of the Triassic. A reticulate, rather than punctuate, model of
speciation is conceived (And. & And. 1983, 1989). We currently recognise
21 species Gondwana-wide (see p. 22).
Dicroidium
UMKOMASIALES
Cj/tRELITZIA 15 (2003)
257
Dicroidium
generic panorama showing
7 of the Molteno species
(see also opposite)
D. elongation
Leaf attachment
Although Dicroidium foliage is so abundant throughout Gondwana,
only one pair of attached leaves (tf. 6a) from the Molteno [And. & And.
1983, pi. 88(1)] and one single attached leaf from Antarctica (Axsmith et
al. 2000) have been reported. A third specimen of importance (tf. 4a) shows
a group of five leaves (and fragments of a further two) arranged in a fasci-
cle but with their base and point of attachment missing [And. & And. 1983,
pi. 88(2)].
Based on the structural similarity of petrified material from the Middle
Triassic Fremouw Fm., Antarctica. Meyer-Berthaud et al. (1993) suggest-
ed that the small stems/twigs they described as Kykloxylon bore Dicroidium
fremouwensis Pigg (1990) leaves. The latter derive from the same locality
but have not been found in organic connection. From the orientation of the
leaf bases they suggest that the ‘distal-most twigs bear helically arranged
leaves’. Considering the Lit 111 specimen (tf. 6a), which shows two leaves
attached at the end of a 40 mm length of slender twig (0.5 mm diam.), and
the group of leaves from Bir 111 (tf. 4a), a terminal leaf fascicle is sug-
gested. This could be accommodated by a close helical phyllotaxy as drawn
in the reconstruction tf. 5.
The single Dicroidium leaf from Antarctica (Axsmith et al. 2000) is
problematic in that it appears to be attached to a robust axis ( ca 15 mm in
diameter) and not to one of the short shoots borne by this same axis. In
extant plants with long and short shoots, the leaves are not directly attached
to mature shoots; e.g. in Ginkgo biloba, leaves occur in fascicles on short
shoots or as individual leaves on long shoots which typically have a thin
axis.
GONDWANA TRIASSIC. GEOSTRAT' DISTRIBUTION
PRE/F/5266b
see And. & And , 1983
pi 88( 1 )
1-10: degn
O: superlo
e squares :
kilties /
2
ir"
a-
3k
’ft
4 ■ ijL
5
A ■ # 6
.
7 ■
>f
C 10
UMKOMASIALES
Dicroidium
Q7 TRELITZIA 15 (2003)
PRE/F/6598b Xl
3P/2/98ia £3? Xl
*■- ■ ■ - r 4.
PRE/F/6602b -s
PRE/F/6602b
a/
'"'I
BP/2/6598bi
BP/2/98 la
Umkomaas Valley
(Umk 1 1 1 Die 2spp)
BP/2/833 "
BP/2/18631
BP/2/842
PRE/F/833
BP/2/18631
NHM London V.23361
V- /
V.23360 NHM London
Holotype
'Warn
BP/2/842 i
Umkomasia macleanii
pi. 82
UMKOMASIALES
^ TRELITZIA 15 (2003)
259
Holotype
PRE/F/8926
Hlatimbe Valley
(Hla 213 Die elo)
PRE/F/8926
UMKOMASIALES
pi. 83
Umkomasia bracteolata
260
c/tRELITZIA 15 (2003)
Umkomasia quadripartita pi. 84 UMKOMASIALES
261
C/ TRELITZIA 15 (2003)
PRE/F/10190a
PRE/F/10190b
PRE/F/9172
PRE/F/9157a
Matatiele
(Mat 1 1 1 Die dub)
BP/2/3172
PRE/F/10190b
PRE/F/2080
PRE/F/9190
PRE/F/10152
PRE/F/9179a
PRE/F/3192
PRE/F/9190
PRE/F/9182
PRE/F/9207
PRE/F/2038
UMKOMASIALES
pi. 85
Umkomasia quadripartita
262
df' TRELITZIA 15 (2003)
U. decussata
Aasvoelberg
(Aas 411 Dic/Sph)
PRE/^1764b^- '
PRE/F/21389a
PRE/F/20827b
PRE/F/20827b
PRE/F/21766
PRE/F/21388b
U. macleanii
PRE/F/4627
PRE/F/4611
PRE/F/4615 D, XI
PRE/F/4608b
PRE/F/4608b
Mazenod
(Maz 21 1 Hei/Dic)
PRE/F/4631
PRE/F/4608b
PRE/F/1688
Umkomasia spp.
pi. 86
UMKOMASIALES
263
c/tRELITZIA 15 (2003)
PRE/F/3168a
PRE/F/2009
PRE/F/2009
PRE/F/9175
Holotype
PRE/F/9173
PRE/F/9174 x2
PRE/F/9174
:'mmr
I PRE/F/9175
PRE/F/9175
PRE/F/9175
PRE/F/2068
PRE/F/9203
PRE/F/2069
UMKOMASIALES
pi. 87
Umkomasia monopartita
264
d^TRELITZIA 15 (2003)
Holotype
PRE/F/5760
BP/2/2100
Little Switzerland
(Lit 111 Dic/Hei)
U mkomasia gracilliaxis
pi. 88
UMKOMASIALES
265
^/tRELITZIA 15 (2003)
Konings Kroon
(Kon 222 Die odo)
U. cupulata
PRE/F/19999
XI 3 PRE/F/19999
BP/2/4122a
Flolotype
PRE/F/20339Jp
BP/2/4122a
(Mor 1 1 1 Die odo)
U. grandis
PRE/F/3396
Holotype
l PRE/F/3397a
UMKOMASIALES
pi. 89
Umkomasia spp.
266
c/trELITZIA 15 (2003)
PRE/F/6653
llmkomaas Valley
(Umk 1 1 1 Die 2spp)
PRE/F/6651 X2
Pteruchus africcinus
pi. 90
UMKOMASIALES
267
TRELITZIA 15 (2003)
UMKOMASIALES
pi. 91
Pteruchus spp.
Hlatimbe Valley
(Hla 213 Die elo)
all Hlatimbe Valley
(Hla 213 Die elo)'
P. matatimajor
^YtrELITZIA 15 (2003)
268
PRE/F/1987a ,xl
PRE/F/1989a
PRE/F/1986a
PRE/F/1988 XI
' ' /'■ V
PRE/F/92 10a
PRE/F/9217
PRE/F/9230a |
PRE/F/9229b
Holotype
Matatiele
(Mat 1 1 1 Die dub)
PRE/F/92 10 >
PRE/F/10149b
PRE/F/10150b
PRE/F/9229b
Pteruch us matatimajor
pi. 92
UMKOMASIALES
269
q/tRELITZIA 15 (2003)
sPRE/F/1989b
PRE/F/1987a
PRE/F/9236
PRE/F/1989a .. l
Matatiele
^ (Mat 1 1 1 Die dub)
PRE/F/10150a
v ;; ^ ' vgJpT''1 - '■
PRE/F/1988
PRE/F/ 1-0 150a
PRE/F/1989b
UMKOMASIALES
pi. 93
Pteruchus matatimajor
270
PRE/F/9225
r "\
PRE/F/9228b
<Sf'' TRELITZIA 15
(2003)
Pteruchus matatimajor
pi. 94
UMKOMASIALES
k I
Matatiele
(Mat 111 Die dub)
PRE/F/9227
A
PRE/F/9227
PRE/F/9228b
PRE/F/9228b -v
X10
271
TRELITZIA 15 (2003)
UMKOMASIALES pi. 95 Pteruchus helvetigracilis
272
cVtRELITZIA 15 (2003)
GINKGOOPSIDA S.V.Meyen 1987
UMKOMASIALES S.V.Meyen 1984
UMKOMASIACEAE S.V.Meyen 1984
F anerotheca Freng. 1944c
Type species
F anerotheca extans Freng. 1944c.
Zanjon. Potrerillos Fm., Cacheuta Basin, Argentina; U. Triassic.
Generic diagnosis emended
A ginkgoopsid strobilus of lax paniculate form with megasporophylls
bearing 1—4 pairs of uni-ovulate cupules with strongly winged seeds.
Generic characters
Strobilus : simple, lax, paniculate, of medium size ( ca 70 mm long); axis
generally robust, erect, gradually tapering; megasporophylls several,
apparently irregularly helical.
Megasporophyll : simple, pedunculate, spicate; bracteoles more or less
prominent, variously scattered along axis and peduncles; ovuliferous
cupules recurved, pedicellate, in 1—4 opposite to subopposite pairs.
Cupule : of medium size (4—8 mm deep), uni-ovulate, enclosing proximal
half of mature seed, splitting into 4 lobes.
Seed: bilaterally symmetrical, strongly platyspermic and strongly winged;
micropyle bifid, not curving to one side.
Etymology
F anerotheca— not given by Frenguelli (1944c).
Global range: 5 spp., Gondwana, Tr. (CRN).
First & last: Molteno Fm.
Gondwana Triassic occurrence
SAm— Cacheuta Basin, 3 or 4 TCs (14 indivs).
SAf— Karoo Basin, 26 TCs (260 indivs).
Aus— Ipswich Basin, 1 TC ( 1 indiv.).
Molteno occurrence (see Tab. 56)
Frequency (F): 26 TCs (of 100 sampled in Molteno).
Diversity (D): 4 species.
Abundance (A): 260 indivs total; rare to very rare in top 6 TCs.
Kap 111 Dic/Ris: 40 indivs in 65 man-hrs cleaving ( 7 per 1 man-day) rare
Wal 111 Die odo: 24 ’
” 50
” ( 4 ”
” ) ”
Bir 111 Hei/Sph: 60 ’
” 550
” (>1 ”
” ) very rare
Lut 311 Hei elo: 7
” 50
” ( 2 ”
” ) ” ”
Umk 1 1 1 Die 2spp: 30
” 400
” ( 1 ”
” ) ” ”
Aas 411 Dic/Sph: 47 1
” 512
” ( 1 ”
” ) ” ”
Affiliated organs
Male strobilus: unknown.
Foliage: Dicroidium— Grade 2 (Mut. occ.).
Classification & comparison
Suprageneric classification (Umkomasiaceae/Umkomasiales)
As Fanerotheca is closely allied to Umkotnasia, we include it in the
family Umkomasiaceae and order Umkomasiales in the class Ginkgoop-
sida.
Intergeneric comparison (Gondwana Triassic)
The lax strobilus of Fanerotheca is in many features like Umkotnasia,
but the cupules with strongly winged seeds (Feruglioa, Frenguelli 1944c)
sets them generically apart.
Townrow (1960) reclassified Fanerotheca as Antevsia, having incor-
rectly interpreted the cupule lobes as sporangial sacs. In And. & And.
[1983, pi. 23(3, 4)], we followed Townrow and used the genus Antevsia for
strobili from Bir 1 1 1 that are here placed in Fanerotheca (F. papilioformis).
The clear attachment of the winged seeds leaves no doubt that this is a
female strobilus and that it is unrelated to Antevsia , a pollen-bearing organ.
Reconstructions
Strobilus
The reconstruction (tf. 1) is based on the type specimen of F. cruci-
formis, PRE/F/10542a,b, pi. 102(1—4), an almost complete strobilus requir-
ing little restoring. The axis may extend basally.
Megasporophyll
A typical megasporophyll has been drawn for each of the four species
(p. 275, tfs 1, 5, 9, 13).
Cupule & seed
Cupules of F. cruciformis with attached winged seeds are shown in lateral
and end view (p. 272, tfs 3, 4). A similar reconstruction has been made for
F. papilioformis and F. waldeckiformis, but not F. elandiformis for which
information on cupules is lacking. All species and localities indicate one
seed per cupule, similar to the case in Umkotnasia.
Fanerotheca
UMKOMASIALES
^/tRELITZIA 15 (2003)
273
Gondwana Triassic occurrence (elaborated)
Fanerotheca was first described from Argentina (Frenguelli 1944c),
where it is fairly common in the Cacheuta area (Tab. 55)— 14 specimens
from three or four localities. Fanerotheca exstans Freng., while lacking
some of our diagnostic features, is retained as the valid type species for
the genus. F. dichotoma (Tab. 55) is here regarded as indeterminate.
Australia has yielded only a single fragmentary specimen, from Denmark
Hill (Walkom 1915), which is the earliest record of Fanerotheca from
Gondwana.
Molteno occurrence (elaborated)
Fanerotheca , occurring at 27 TCs (Tab. 56), is the most frequent of all
ovulate genera in the Molteno. The next most frequent is Umkomasia,
which occurs at 22 TCs (Tab. 52). However, the reverse holds when con-
sidering abundance, Umkomasia with 503 individuals being more abundant
than Fanerotheca with 247 individuals.
Tab. 55
FANEROTHECA HYPODIGM, Gondwana Triassic occurrence
Species
Intact-
ness
Molteno
F. papilioformis
F. watdeckiformis
F. cruciformis
F. elandiformis
F. exstans
F. spp. indet
Intact strobili
Fragmentary
Isolated megasp.
AUTHOR
SUBREGION (°sq.)
FORMATION
LOCALITY
NAME (original)
Indivs ILLUSTRATION
SOUTH AMERIC/
1
1 1 1
1 1 1
1 1 1
1
1
1 ' 1
1 1 •
1944c! Frenguelli
Cacheuta NA4
24 Potrerillos
Zanjon
Fanerotheca exstans
4 : pi 1(1,2), 2(1), 3(1,2)
4!-
212,-
» 1 i>
1
1
YPF Old Admin. Bid.
3 pi 3(3), 4(1,2)
i _ ! _ 1 _
3 : -
3
1
” |
?Locality
" dichotoma
1 pi 2(2,3)
- i - i . i -
- ! i
- ; 1 : -
1946 i
” | - ”
-
Stachyopitys anthoides
1 pi 4(3)
- 1 - 1 ~ 1 ~
1 1 -
1 1 - 1 -
1960 Townrow (specimen repeated from Frenguelli 1944c, pi 1(1))
Antevsia exstans
* 1 -
-!-!-!-
* i -
* i - 1 -
1967 ! Jain & Del.
” | ”
>. 1 ”
Minas de Petroleo
Fanerotheca exstans
3 pi 90(11-13)
3 ! -
- 1 1 2
” ;
„ i »
„ i
incertae sedis
2 pi 90(9,10)
i i i
i i i
2T7
-!2 ;-
SOUTH AFRICA
!
1
i
l i I
l i l
I
1
1927 DuToit
Elliot Ka9
24! Molteno
Cala (2mls West)
Sagenopteris sp.
1 pi 29(3)
- i - i - i -
- 1
- 1 - 1 1
1960 ! Townrow
Underberg Ka4
24 ■
Upper Umkomaas
Antevsia exstans
1 tf 7(E.F), pi 58(6)
- 1
- 1 - 1 1
1979-1999 And. & And. Molteno literature not included in this table
I l l
1
AUSTRALIA
1
1
1
1
1 l I
1 I l
1
1
* 1
1915 Walkom
Ipswich/Esk <CM5
24 Blackstone Stage
!
Denmark Hill
Equitsetites sp.
1 pi 3(3,4)
L—J 1
- 1 1
-;ii-
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
UMKOMASIALES
Fanerotheca
274
c/tRELITZIA 15 (2003)
F. papitioformis
tfs 1,2
PRE/F/18694
prep. no. 1040b
from cupule lobe
Evidence for afniiation of organs
Foliage
There is no clear pattern of mutual occurrence for Fanerotheca and
any foliage affiliate.
Fanerotheca is particularly common at Bir 111 and Aas 41 1, where it
seems to have a link with the abundant Sphenobaiera (Tab. 50). But
Sphenobaiera has now been securely affiliated with Hamshawvia and the
attached immature fertile organ is definitely not Fanerotheca. Furthermore,
at Kap 111 where Fanerotheca is also common, there are only 20 speci-
mens of Sphenobaiera (under 1%).
At Bir 111 and Wal 111, although the frequency of Dicroidium is 1%
and 92% respectively, no Umkomasia has been found (Tab. 56). This has
led to our suggestion that Fanerotheca may be linked to Dicroidium, which
is supported by: ( 1 ) the similarity in strobili and cupule structure of
Fanerotheca and Umkomasia, and (2) Fanerotheca occurring in 13 TCs
with Dicroidium but no Umkomasia (Tab. 52). However, at these 13 TCs
there is no clear correlation with any particular Dicroidium species or
group of species.
Male strobilus
The male affiliate is unknown, but it could possibly be a species of
Pteruclms or Stachyopitys (see Tabs 50 and 54 for the distribution of these
two genera along with that of Fanerotheca).
Seed
The seed now known attached to Fanerotheca (Grade 5 affiliation)
was originally described by Frenguelli (1944c) as Fertiglioa samaroides.
From the same locality, Frenguelli also described Fanerotheca extans, but
he did not record any of the seeds as being attached.
Intactness & preservation
In situ seeds
F. cruciformis— In the most complete strobilus from Bir 111,
PRE/F/10542a,b, pi. 102(1^1), four seeds, one per cupule, remain attached.
These appear to be mature and are ca twice as long as the cupule.
F. papilioformis— At Bir 111, one strobilus shows a single attached
seed which is twice the length of the cupule lobes [BP/2/5 199b, pi. 97(2,
3)]. Another shows several seeds attached, one to a cupule [BP/2/11854, pi.
96(3, 12)]. Further specimens show attached seeds but these are not illus-
trated here: Bir 111, BP/2/5201a, BP/2/5007; Umk 111, PRE/F/1694.
F. waldeckiformis— At Wal 111 all specimens have lost their seeds,
while at Mat 111a single strobilus, PRE/F/2010, pi. 101(8), shows seeds
still attached to the cupules.
Germinating seeds
At Bir 111, the winged seeds, Feruglioa, occur massed on certain bed-
ding planes. Some show evidence of having germinated by the presence of
a distinct radicle emerging from the micropyle end of the seed. This can be
seen in various specimens, some also with their first leaves (tfs 3-7 adja-
cent; pi. 99). Krassilov (1987, pi. 4) illustrates very similar germinating
seeds and seedlings, with clear elongate leaves, from the Jurassic of Ust-
Balej (USSR).
Cuticles
Potential sample'. Lit 1 1 1, 3 indivs; Umk 111, 16 indivs.
Macerated (this work): Umk 111,2 indivs from cupule lobe.
Preser\’ation grade : Grade 3, some features including stomata visible.
Diagnostic characters: cells isodiametric to narrowly pentagonal to linear-
oblong, walls gently curved, nonpapillate; stomata (?)amphistomatic,
orientation longitudo-random, subsidiary cells nonlappetate brachy-
paracytic, guard cells narrowly elliptic.
Comment: the illustrated cuticle derives from the cupule lobe.
Significance:
Classification— Though the macerated cuticle is thin and not too clear-
ly preserved, it is sufficient to support the classification of Fanerotheca in
the order Umkomasiales and class Ginkgoopsida.
Affiliations— In its subsidiary cells being nonlappetate and brachy-
paracytic, Fanerotheca is clearly more like Dicroidium than any of the
other typically ginkgoopsid foliage genera such as Lepidopteris,
Sphenobaiera, Ginkgoites and Dejerseya. In being nonpapillate and having
narrowly elliptic guard cells, it most nearly approaches the cuticle of D.
odontopteroides forma odontopteroides (And. & And. 1983, p. 199).
However, it does not show the cutinised subsidiary cells or trichome bases
of the latter taxon.
Fanerotheca
UMKOMASIALES
'^/tRELITZIA 15 (2003)
275
Adaptive radiation (Molteno diversity)
Although Fanerotheca occurs frequently and commonly in the
Molteno— 247 individuals (65 of which are intact) from 27 TCs— it is
markedly lacking in evident diversity. Only four species can be differenti-
ated. The diagnostic characters lie essentially in the robustness of the stro-
bilus, the number of cupule pairs, and in the nature of the cupules and
length of their pedicels.
The four species, based on very variably sized palaeodemes, are all
from floodplain habitats— either Dicroidium or Sphenobaiera woodland.
This is in marked contrast to the Umkomasia species which are mainly
from Dicroidium forest habitats.
F. papilioformis —B'\r 111 Sph 2 spp (Birds River), 52 indivs
Sphenobaiera closed woodland (floodplain lake); Cycle 5 (Tsomo Member)
F. waldeckiformis—Wal 111 Die odo (Waldeck), 24 indivs
Sphenobaiera closed woodland (floodplain lake); Cycle 2 or 3
F. cruciformis— Bir 111 Sph 2 spp (Birds River), 8 indivs
Sphenobaiera closed woodland (floodplain lake); Cycle 5 (Tsomo Member)
F. elandiformis — Ela 112 Dic/Hei (Elandspruit), 2 indivs
Dicroidium open woodland (floodplain); Cycle 2a (Indwe Member)
Tab. 56. Fanerotheca, Molteno occurrence
UMKOMASIALES
Fanerotheca
276
ry TRELITZIA 15 (2003)
F anerotheca papiliofonnis J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/ 10544a, b; pi. 96(1, 13).
Assemblage (TC): Bir 1 1 1 Sph 2spp, Birds River.
Presen’ation: complete specimen, part and counterpart; impression in thinly
laminated, yellowish grey shale with very good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 52 indivs (21 intact, 23 partial, 8 isolated), pis 96-99.
Sister palaeodemes— 15 (best 2 listed)
Aas 411 Dic/Sph: 47 indivs (11 intact, 26 partial, 10 isolated).
Kap 111 Dic/Ris: 40 indivs (3 intact, 26 partial, 11 isolated).
Specific diagnosis
A Fanerotheca species with gracile strobili bearing megasporophylls
with 1 or 2 pairs of cupules on short pedicels.
Specific characters
Megasporophyll: gracile; cupules in 1 or 2 pairs, opposite and decussate;
pedicels short (1-3 mm).
Cupule: gracile, 2- to 4-lobed, with lobes deeply divided to pedicel attach-
ment and folding tightly along midline.
Ovule! seed: relatively small (7.5-11 mm long), body occasionally covered
with characteristic oval spots.
Etymology
papiliofonnis— papilio (Lat.), butterfly, with reference to the butterfly-
shaped cupule lobes.
Comments & comparison
This is the most common and widespread (17 TCs) of the Molteno
Fanerotheca species. It is very distinctive in that the lobes are always
deeply divided to the point of attachment with the pedicel. The spots on the
seed body may be secondary, i.e. of fungal origin.
Fanerotheca waldeckiformis J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/9440a,b; pi. 100(2—4).
Assemblage (TC); Wal 111 Die odo, Waldeck.
Preservation: almost complete specimen, part and counterpart; impression
in thickly laminated, moderately baked, medium light grey shale with very
good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 24 indivs (10 intact), pis 100(1-10), 101(1-7).
Sister palaeodemes— 4 (best 1 listed)
Matlll Die dub: 6 indivs (2 intact, 3 partial, 1 isolated), pi. 101(8-13).
Specific diagnosis
A Fanerotheca species with fairly gracile strobili bearing megasporo-
phylls with 1 or 2 pairs of cupules on long pedicels.
Specific characters
Megasporophyll: gracile; cupules in 1 or 2 pairs, clustered distally; pedicels
long (4—8 mm).
Cupule: fairly robust, 4-lobed, with lobes divided ca midway to pedicel attach-
ment.
Ovule/seed: intermediate in size (10-14 mm long), body without spots.
Etymology
waldeckiformis— with reference to the type locality.
Comments & comparison
This species is distinguished from F. papiliofonnis in the longer
pedicels and shallower cupule lobing, and from F. cruciformis in the fewer
cupule pairs.
F anerotheca
UMKOMASIALES
Cff TREL1TZI A 15 (2003)
277
Fanerotheca cruciformis J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/10542a,b; pi. 102(1-4).
Assemblage (TC): Bir 1 1 1 Sph 2spp, Birds River.
Preservation: almost complete specimen, part and counterpart; impression
in thinly laminated, yellowish grey shale with very good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 8 indivs (1 intact, 1 partial, 6 isolated), pi. 102(1-7).
Sister palaeodemes— 1 only (as listed)
Maz 1 11 Die era: 6 indivs (1 intact, 2 partial, 3 isolated).
Specific diagnosis
A Fanerotheca species with robust strobili bearing megasporophylls
with 3 or 4 pairs of cupules on short pedicels.
Specific characters
Megasporophyll: robust; cupules in 3 or 4 pairs, subopposite and decus-
sate; pedicels short (1-3 mm).
Cupule: fairly robust, 4-lobed, with lobes divided ca 3/4 to pedicel attach-
ment.
Ovule/seed: intermediate in size (10-12 mm long), body without spots.
Etymology
cruciformis— with reference to the cross-shaped cupule lobes when open.
Comments & comparison
This second species of Fanerotheca from Bir 111, with only eight indi-
viduals (one intact), is by far the less common at this site. It is distinct from
other species in being more robust and bearing three to four pairs of
cupules.
Fanerotheca elandiformis J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/13306a,b; pi. 103(6,7).
Assemblage (TC): Ela 112 Dic/Hei, Elandspruit.
Preserx’ation: incomplete specimen, part and counterpart; impression in
thickly laminated, medium grey shale with poor cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 2 indivs (2 partial), pi. 103(1-9).
Sister palaeodemes— nil.
Specific diagnosis
A Fanerotheca species with strobili bearing megasporophylls with (?)2
or 3 pairs of naked seeds on fairly long pedicels.
Specific characters
Megasporophyll: robust; naked seeds in (?)2 or 3 pairs; pedicels of inter-
mediate length (3-5 mm).
Cupule: unknown.
Ovule/seed: relatively large (11-16 mm long), body invariably covered
with characteristic protuberances.
Etymology
elandiformis— with reference to the type locality. Elandspruit.
Comments & comparison
In view of the large seeds, this is clearly a distinct species. Also, the
characteristic four-lobed cupules found in the other species are apparently
absent— as witnessed in the holotype, the only partly intact specimen,
which has naked seeds attached directly to the pedicels [pi. 103(7)].
The unique markings on the seeds are generally spherical, protruding,
and differ from the more level, oval variety characterising F. papilioformis.
It is possible that these markings are a secondary feature, such as fungal
spots, in which case they are apparently host-specific. In F. elandiformis
they occur consistently on all collected seeds, not the case in F. papilio-
formis.
UMKOMASIALES
Fanerotheca
278
c/trELITZIA 15 (2003)
PRE/F/11854
BP/2/5203
BP/2/5202
PRE/F/10544a ■;
Holotype
PRE/F/10021 j
PRE/F/10818
PRE/F/15585 X2
PRE/F/15615b
PRE/F/15585
PRE/F/10021
Birds River
(Bir 111 Sph 2spp)
BP/2/5203
PRE/F/11854
PRE/F/1 0544a
Fanerotheca papilioformis
pi. 96
UMKOMASIALES
279
d/TRELITZIA 15 (2003)
all BP/2/5199b
Birds River
(Bir 1 1 1 Sph 2spp)
pi. 97
F anerotheca papilioformis
UMKOMASIALES
TRELITZIA 15 (2003)
PRE/F/1 1786a
BP/2/5356
PRE/F/10800
JPRE/F/10082
Birds River
(Bir 1 1 1 Sph 2spp)
PRE/F/10082
BP/2/5352
Fanerotheca papilioformis
pi. 98
UMKOMASIALES
281
^I/tRELITZIA 15 (2003)
PRE/F/1 1791b
BP/2/5365
BP/2/5336
BP/2/5376
BP/2/5394
BP/2/5374
BP/2/5376
BP/2/5368
Birds River
(Bir 1 1 1 Sph 2spp)
r BP/2/5368
PRE/F/10075
UMKOMASIALES
pi. 99
Fanerotheca papilioformis
TRELITZIA 15 (2003)
PRE/F/9442
Waldeck
(Wal 1 1 1 Die odo)
PRE/F/9440a
Flolotype
PRE/F/9440a
PRE/F/9535b
PRE/F/9440a
PRE/F/9535b
PRE/F/9538a
PRE/F/9538b
F anerotheca waldeckiformis
pi. 100
UMKOMASIALES
283
c/tRELITZIA 15 (2003)
Matatiele
(Mat 1 1 1 Die dub)
PRE/F/9440a'y'
PRE/F/9455
PRE/F/2010
BP/2/3173
PRE/F/9536
PRE/F/9537
Waldeck
(Wal 1 1 1 Die odo)
PRE/F/9536b
pi. 101
F anerotheca waldeckiformis
UMKOMASIALES
284
d/TRELITZIA 15 (2003)
PRE/F/10542b
PRE/F/10542a
Birds River
(Bir 111 Sph 2spp)
PRE/F/10542b
Holotype
m
PRE/F/10816
BP/2/5210
PRE/F/10542a
Fanerotheca cruciformis
pi. 102
UMKOMASIALES
UMKOMASIALES
pi. 103
F anerotheca elandiformis
286
d^TRELITZIA 15 (2003)
GINKGOOPSIDA S.V.Meyen 1987
PETRIELLALES T.N.Taylor etal. 1994
KANNASKOPPIACEAE J.M.And. & H.M.And.. fam. nov.
Kannaskoppia J.M.And. & H.M.And., gen. nov.
Type species
Kannaskoppia vincularis J.M.And. & H.M.And., sp. nov.
Kannaskop, Karoo Basin, S. Africa; Carnian, Triassic.
Generic diagnosis
A ginkgoopsid strobilus with a forked axis bearing rows of several
simple megasporophylls consisting of a single cupule.
Generic characters
Fertile shoot', stem with short shoots in an irregular helical arrangement;
leaves (1 or 2) and strobili (2 or 3) borne irregularly on each shoot.
Strobilus : lax, planar, bilaterally symmetrical, small (ca 20 mm long); axis
gracile, proximally forked; megasporophylls in 2 adaxial oblique rows
(of 4-6 units) along each fork (limb), angle between rows ca 90°.
Megasporophyll: apparently reduced to single pedunculate cupules; ovulif-
erous cupules recurved; peduncles 2-3 mm long, gracile, sinuously
curved.
Cupule'. small {ca 2.5 X 2 mm), roundly ovoid, splitting more or less regu-
larly into 3 lobes at maturity.
Ovule/seed: unknown.
Etymology
Kannaskoppia— after the type locality Kannaskop, an Afrikaans name for a
hill bearing Canna plants (monocots of the family Cannaceae).
Global range: 1 sp.. Gondwana, Tr. (CRN).
First & last: Molteno Fm.
Gondwana Triassic occurrence
SAf— Karoo Basin, 1 TC (50 indivs).
Molteno occurrence (see Tab. 57)
Frequency (F): 1 TC (of 100 sampled in the Molteno).
Diversity (D): 1 species.
Abundance (A): 50 indivs; rare.
Kan 1 1 1 Ast spA: 50 indivs in 30 man-hrs cleaving (ca 20 per 1 man-day), rare
Affiliated organs
Male strobilus : Kannaskoppianthus— Grade 5 (Mor. corr.. Kin. reinf.).
Foliage : Kannaskoppifolia— Grade 5 (Org. att. ).
Classification & comparison
Suprageneric classification (Kannaskoppiaceae/Petriellales)
Kannaskoppia is considered here to be closely allied to Petriellaea and
thus placed in the order Petriellales (Taylor et al. 1994). However, in view
of the uncertainties that remain (see below), we include the Molteno genus
in a distinct family, the Kannaskoppiaceae.
Intergeneric comparison (Gondwana Triassic)
Petriellaea is known from a single permineralised peat deposit in the
Middle Triassic Fremouw Fm. of Antarctica (Taylor et al. 1994). The
microscopic anatomy of the ovulate organ is exquisitely preserved reveal-
ing five ovules per cupule and many other details, but the overall architec-
ture of the fruit remains uncertain. The reflexed cupules, as well as the size
and configuration of the ovulate heads, are reminiscent of Kannaskoppia.
It is possible that further new finds may show Petriellaea and
Kannaskoppia to be congeneric, but this is not possible to conclude at the
present time. What is known of the one genus is mostly unknown of the
other.
Reconstructions
Fertile shoot
The reconstructions (tfs 1-3 above; tfs 1, 2, p. 288) are based on the
comprehensive palaeodeme from Kan 1 1 1 Ast spA. The shoot showing
attachment of foliage and female fruit (p. 288, tf. 1) is largely a combina-
tion of PRE/F/13487a‘y' and PRE/F/13487b‘x’— the two portions of shoot
illustrated on p. 289 (tfs 1 , 2). These lie closely adjacent on the part and
counterpart of a single cleaved slab. The arrangement of leaves and megas-
porophylls on the short shoots appears to show no consistent systematic
pattern. However, the size and number of megasporophylls probably
diminish towards the shoot apex.
Strobilus
The strobili, occurring in twos or threes, are found attached to bulbous
short shoots which usually bear one or two leaves. Simplified reconstruc-
tions, with leaves removed, include a group of three strobili at natural size
(p. 288, tf. 2), and a single strobilus showing characteristic fork and double
row of megasporophylls at x2 magnification (tf. I above).
Megasporophyl l
The megasporophyll is taken to consist of a single cupulate unit borne
on a long gracile, slightly recurved peduncle. Cupule details (tfs 3a-d above)
are based largely on three specimens, PRE/F/ 1 3489a,b, PRE/F/13508a,b and
PRE/F/13518a,b; see pi. 105(1-3, 4), which show the cupules preserved at
various angles. Due to the 3D nature of the preservation of the material
shown in pi. 105(4), it is possible to observe the curved peduncle with its
tendency to flex downwards, and to estimate the angle of ca 90° between
the cupule rows as shown in our reconstructions (tfs 1, 2 above; tfs 1, 2, p.
288). The cupule is roundly ovoid and at maturity splits into three more or
less regular lobes (tfs 3c-d above).
Ovule/seed
While much is known about the gross morphology of the megasporo-
phylls and their mode of attachment, nothing is known of the interior
organisation of the cupules or of the ovules. Nowhere on the many slabs
bearing the fruit is there any indication that either cupules or seeds are indi-
vidually dispersed.
Kannaskoppia
PETRIELLALES
d/TRELITZIA 15 (2003)
287
assemblages
(taphocoenosis)
Kannaskoppifolia
O Kannaskoppia
Q. Kannaskopianthus
microsporangia
man-hours
cleaving
fruit abundance:
indivs./man-day
O, K. lutinumerus
" K. matatiparvus
" K. irregularis
” K. telemagnus
Cal 211 Hei elo
i i i
5 - - -
1
- I
i i i
- , - i - i -
Gre 121 Hei elo
22 1 - 1 2 1
10 1/1
2! - ! - [ -
” 111 Equ sp
i ! -
25
1 j ; -|
Boe 112 Die cor
i ! -
1
_ 1 _ 1 . t .
Cyp 111 Die era
83 -; -
100
. . . : . i .
Kan 112 Hei elo
19 - 5 1
15 ' 3/1
5 - 1 5 ! -
” 111 AstspA
5 1 50 - 1 -
30 20/1
Tel 111 Hei elo
33 ! - 41
90 1/2
2! - !_J 2
Korn 111 Sph/Dic
30 - 2 ; 1
10 2/1
-! - ! 2 ! -
Vin 111 Die odo
2; -: -
- 1 - 1 - 1 -
Lut 311 Hei elo
66; -! 16 1 -
50 > 3/1
16 - ! - ! -
Kon 211 Ast 2spp
4 1 -> -i -
- 1
- , “ 1 - 1 -
Pen 311 Hei elo
4i -! - ! -
35 !
“ - 1 - -
„ 411 .. »
70 - 4 -
70 1/2
4 _ 1 _ 1 _
Kap 111 Die spp
6 ! -! -! -
65 -
-l . 1 - 1 -
Nuw 111 Die zub
i .) .
21 1
- 1 - 1 - ! -
Win 111 Hei elo
4! -i -i -
20
_ 1 _ 1 _ 1
Hla 213 Die elo
7 — ! -! -
60
_ 1 . 1 _ 1 _
Umk 111 Die 2spp
42 - 1 - i -
400 . -
. 1 . « . 1
San 111 Die era
3 -; -
30
- ! - ! - 1 -
Mat 111 Die dub ■
2 u 3 -
65 1/2
- ; 3 ! - < -
Lit 111 Dic/Hei
56 - 1 9 > 2
550 1/6
9 - 1 - 1 -
Aas 111 Hei elo
2 - 21 5
40 5/1
21 - - -
„ 211 „ „
19 - 1 -
35 1/3
1 1 - ; - ; .
„ 311 ..
20 1 -! 4\ 1
140 1/3
4 . j . ; .
" 411 Dic/Sph
150 \ - ; 21 ; 4
512 | 1/3
21 - - 1 - 1 -
Total TCs
26| 1 ! 12 8
!
10 1:21
Total individuals
% ; 50 ! 92 ; 16
1
80 3 7 2
Tab. 57. KannaskoppialKannaskoppifolia,
Molteno occurrence
Evidence for affiliation of organs
Organic attachment
The holotype specimen of Kannaskoppifolia vincularis, together with
the associated specimen on the same slab and additional material, consti-
tutes the most convincing example of affiliated organs in the Molteno flora.
Typical strobili and leaves are found attached to short shoots borne on a
section of stem. Other specimens in the collection show the male strobilus,
Kannaskoppianthus, attached in a similar manner to short shoots with
Kannaskoppifolia leaves.
Comparisons beyond Gondwana Triassic
Apart from Petriellaea , the Mesozoic ovulate structure most
comparable with Kannaskoppia is Caytonia (Jurassic, Eurasia)— in
its size and form with lateral rows of dorsally reflexed cupules on
short peduncles. In both genera the cupules fully enclose the ovule(s),
but in Kannaskoppia their nature remains unknown. Kannaskoppia
differs most evidently in the strobilus being forked and in the pedun-
culate cupules not abscising.
Caytonia (Jurassic/Eurasia)
Geographic <£ stratigraphic distribution : The ovulate organ, Caytonia ,
is now known as a widespread element of the lower half of the
Jurassic of Eurasia: Yorkshire (>9 localities, M. Jurassic, Bajocian to
Bathonian); Greenland (2 localities, basal Liassic, Hettangian);
Poland (1 locality, U. Liassic); Sardinia and the USSR. Foliage iden-
tified as Sagenopteris has been reported from the U. Triassic to U.
Cretaceous. More recently, the first report of Caytonia was made
from Gondwana. Clifford & Camilleri (1998) describe C. tierneyi
affiliated with Sagenopteris leaves from the Lower Jurassic Marburg
Fm., Queensland, Australia.
Affiliation of organs: The Caytonia plant, the basis of the order Cayton-
iales, is one of around 20 fossil and extant gymnospermous 'orders’
(or approximate ordinal groupings) on which recent cladistic analy-
ses of the gymnosperms are based (Crane 1985, 1986, 1988). It is one
of the few fossil gymnosperm genera where most organs are known
through well-established affiliations (Grade 4) from a good number
of localities. The organs include Caytonia (female strobilus),
Caytonanthus (male strobilus) and Sagenopteris (foliage).
Preservation : Adding value to Caytonia (and affiliated organs) is that
it is often found as well-preserved compressions.
Megasporophylls (morphology): ‘Cupules’ on short peduncles, spher-
ical, reflexed dorsally with distinctive lip, apparently fleshy and berry-
like at maturity, abscising at base of pedicel, with 8-30 tiny ovules.
Abundance: Explicit abundance and frequency data on fossil taxa are
rarely given and this holds generally true for the Caytonia plant. In
the Yorkshire Jurassic sites where it is best known, Harris (1964, p. 3)
writes that it is ‘by no means common’.
Diversity: No recent taxonomic revision of the Caytonia plant has
been attempted, so any real sense of specific diversity is very difficult
to gather. Five named species of the genus Caytonia appear to be cur-
rently valid.
References: Harris (1933, 1940, 1958, 1964); Reymanowna (1973);
Crane (1985); Krassilov (1977).
Polyspermophyllum ( Permian/ Argentina)
Kannaskoppia cupules are remarkably similar to the ovuliferous
component of Polyspermophyllum sergii Archangelsky & Cuneo
(1990) from the Permian of Argentina. These are described as ‘dis-
tally placed curved ovuliferous units (cupules?) bearing a single
ovule.’ However, the ?cupules are borne on repeatedly dichotomising
trusses and occur with attached leaves bearing a single medial vein
and two longitudinal marginal furrows, setting them far apart from
Kannaskoppia. Polyspermophyllum is included in the Dicranophyl-
lales. Archangelsky & Cuneo also discuss the possible links of their
genus to the Ginkgoales and other orders.
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
PETRIELLALES
Kannaskoppia
Kannaskoppia
PETRIELLALES
& : TRELITZIA 15 (2003)
289
Kannaskoppia vincularis j.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/ 13487 a‘y’; tf. 2 below; pi. 104(3, 5).
Assemblage: Kan 1 1 1 Ast spA, Kannaskop.
Preservation: a shoot 90 mm long with clusters of foliage and megasporo-
phylls attached; compressed 3D mould and cast (no cuticle or carbonised
material); on the counterpart slab there is an additional isotype specimen,
PRE/F/ 1 3487b ‘x\ from which a piece of matrix can be lifted to reveal
foliage and fruit at a different level; impression in thick-bedded, moder-
ately baked, greenish grey silty mudstone with very poor cleavage.
Reference palaeodeme
Assemblage: as for holotype.
Specimens: ca 50 indivs, pis 104, 105 (includes further foliage as indicated);
5 tom sections of shoot with foliage and strobili attached;
>50 detached strobili or clusters of strobili;
12 tom sections of shoot with foliage only;
>25 detached leaves, generally tom or twisted.
Sister palaeodemes— nil.
Specific diagnosis— as for genus.
Specific characters— as for genus.
Etymology
vincularis— vinculum (Lat.), link, bond, with reference to the fruit and foliage
being found in organic connection at the type locality.
Comment & comparison— as for genus.
K. vincularis
PETRIELLALES
Kannaskoppia
290
G7TRELITZIA 15 (2003)
Kannaskoppianthus J.M.And. & H.M.And., gen. nov.
Type species
Kannaskoppianthus lutinumerus J.M.And. & H.M.And., sp. nov.
Lutherskop, Karoo Basin, S. Africa; Carnian, Triassic.
Generic diagnosis
A ginkgoopsid strobilus comprising a forked axis bearing rows of several
simple microsporophylls, each with 5 longitudinal microsporangia in a con-
cavity protected by an operculum.
Generic characters
Fertile shoot : strobili attached in fascicle with leaf cluster along shoot or at
end of shoot.
Strobilus : lax, planar, bilaterally symmetrical, small (8-25 mm long) to rarely
medium (up to 45 mm); axis gracile, proxirrially forked; microsporo-
phylls in 2 adaxial oblique rows (of 8-10 units) along each fork (limb),
angle between rows ca 90°.
MicrosporophyU : spathulate, dorsiventrally flattened, bilaterally symmetri-
cal; microsporangia 5 per unit, longitudinally aligned in distinctive
concavity, attached distally to apical arcuate scale cap, protected by a
dehiscent operculum.
Microsporangia: elongate-elliptic (1.5 x 0.5 mm), with fine linear ornamen-
tation.
Pollen: unknown.
Etymology
Kannaskoppianthus —After the type locality and acknowledging certain affili-
ation with the female strobilus Kannaskoppia.
Global range: 4 spp., Gondwana, Tr. (CRN).
First & last: Molteno Fm.
Gondwana Triassic occurrence
SAf— Karoo Basin, 12 TCs (92 indivs).
Molteno occurrence
Frequency (F): 12 TCs (of 100 sampled in Molteno).
Diversity (D): 4 species.
Abundance (A); 92 indivs total; rare to extremely rare in top 8 TCs.
Aas 111 Hei elo: 21 indivs in 40 man-hrs (5 per 1 man-day) rare
Lut 311 Hei elo:
Kan 112 Hei elo:
Kom 111 Sph/Dic:
16 ”
5 ”
2 ”
” 50
” 15
” 10
” (3
” (3
” (2
’ 1
‘ 1
‘ 1
” ) ”
” ) ”
” ) ”
Aas 411 Dic/Sph:
21 ”
” 512
” (1
’ 2
” ) very rare
Mat 111 Die dub:
3 ”
” 65
” (1
’ 2
” ) ” ”
Tel 111 Hei elo:
4 ”
” 90
” (1
’ 2
” ) ” ”
Lit 111 Dic/Hei:
9 ”
” 550
” (1
' 6
” ) extr. rare
The abundance figures reflected here for the eight TCs with the high-
est Kannaskoppianthus yield, account for every specimen found, no matter
how fragmentary or poorly preserved. All were retained and curated. At the
four further sites not listed above, Kannaskoppianthus is even rarer (Tabs
57, 58).
Although very rare, the Kannaskoppianthus male occurs in 12 of 25
TCs yielding Kannaskoppifolia foliage. This frequency ratio of 1:2 (male
to foliage) is high for the Molteno. In the 12 TCs it ranges in abundance
from 1 individual in 7 man-days cleaving to 5 in 1 day, with a norm at
around 1 found every 2 days. There appears to be no particular pattern of
abundance based on habitat.
Affiliated organs
Female strobilus: Kannaskoppia— Grade 5 (Mor. corn).
Foliage: Kannaskoppifolia— Grade 5 (Org. att. ).
From Kommandantskop (Kom 111 Sph/Dic), PRE/F3231a,b (tf. 5
adjacent) shows two strobili of Kannaskoppianthus attached to an axis with
a whorl of four Kannaskoppifolia leaves (see also tf. 2, p. 293). From
Kannaskop (Kan 112 Hei elo), PRE/F/201 14a,b (tf. 1, p. 293) shows a stro-
bilus attached to a stem with leaf/branch scars. This is similar to the stems
bearing leaves and female strobili ( Kannaskoppia ) from Kan 1 1 1 Ast. sp.A.
Classification & comparison
Suprageneric classification
See under Kannaskoppia.
Intergeneric comparison (Gondwana Triassic)
Kannaskoppianthus is unique among known ginkgoopsid microspo-
rangiate genera. In their size, diagnostic forking and paired erect rows of
fertile units, the micro- and megasporophylls ( Kannaskoppia ) are remark-
ably alike. The latter are noticeably more gracile.
Kannaskoppianthus
PETRIELLALES
d/TRELITZI A 15 (2003)
291
Reconstructions
The R5 reconstructions (tfs 1, 2 opposite) of Kannaskoppianthus are
based on the full set of Molteno specimens at hand, but primarily on:
Lut311; PRE/F/1 1433, pi. 110(3,7): architecture of strobilus, nature of protective
operculum
Aas211; PRE/F/15280, pi. 113(1—4-): angle of microsporophyll
Tel 111; PRE/F/18277a,b. pi. 116: structure of microsporophyll, attachment and number
of microsporangia
Aas411; PRE/F/12050 & 21720. pi. 115: attachment of microsporangia
Aas311; PRE/F/1 200 1, p. 290, tf. 4, pi. 114(1-3): morphology of microsporangia
Tel 111; PRE/F/77 1 1 , pi. 118: nature of protective operculum
Lit 111; PRE/F/21497: number of microsporangia, groove (?vascular strand)
Gre 121; PRE/F/7856: number of microsporangia, groove (?vascular strand)
Number of microsporangia
No specimens in the collection show microsporophylls with the full
complement of five microsporangia attached. A specimen from Tel 111,
PRE/F/1 8277a,b, clearly shows two attached microsporangia, pi. 1 16(1-9).
The size and shape of these microsporangia suggest a full complement of
five pollen sacs in the undehisced original. Furthermore, five grooves or
vascular strands are seen where the microsporangia were probably attached
(tf. 2d opposite).
Microsporophyll developmental & dehiscence stages (tfs 2 a-e opposite):
(a) microsporophyll with protective operculum covering the microsporangia;
(b) operculum dehisced revealing the 5 closely packed immature micro-
sporangia;
(c) mature microsporangia ready to release pollen;
(d) microsporophyll with the microsporangia detached exposing 5 grooves
or vascular strands;
(e) the woody arcuate scale cap is now detached; most specimens in the
collection show this stage of preservation;
(f) microsporophyll at stage (b) in oblique view at characteristic angle (as
if on intact strobilus).
Gondwana Triassic occurrence (elaborated)
A doubtful record of a sporophyll was reported by Jones & De Jersey
(1947, tf. 59) from the Ipswich Basin in Australia. Although thought to bear
seeds, in size and shape the specimen comes close to Kannaskoppianthus.
However, poor preservation does not allow closer comparison.
Comparisons beyond Gondwana Triassic
The only pollen-bearing structures that show some similarity to
Kannaskoppianthus occur in the Carboniferous of Euramerica. The
class Lagenostomopsida includes the genera Crossotheca and
Feraxotheca. both of which have numerous microsporangia attached
at the end of modified ultimate pinnae. Paracalathiops shows a sim-
ilarity in the basic architecture of a bifurcating axis and pedunculate
microsporophylls (see Taylor & Taylor 1993).
While the megasporophyll Kannaskoppia shows some similari-
ties to Caytonia, the microsporophyll Kannaskoppianthus differs
greatly from Caytonianthus , which has a pinnate structure bearing
lateral ‘branches’ and a few pedicellate microsporangia consisting of
usually ‘four locules’ which dehisce towards their inner side (Crane
1985).
Intactness & preservation
Microsporangia (see Tab. 58)
The male strobilus has been found in 12 of the 25 Kannaskoppifolia-
bearing TCs. Although microsporangia have been recognised from eight
of these 1 2 TCs they remain notably rare, probably because the micro-
sporangia are shed before or during fossilisation. Of the strobili preserved
with associated microsporangia, only a small proportion are seen with sacs
clearly attached: Tel 111 (PRE/F/1 8277a, b); Aas 411 (PRE/F/1 2050a,b,
PRE/F/20558a,b, PRE/F/21339a,b, PRE/F/21720a,b). The best preserved
of the individual microsporangia have been illustrated: pi. 114(1-5) from
Aas 31 1, pi. 115(7) from Aas 411, and pi. 1 16(1-9) from Tel 111.
Pollen
Of all the Kannaskoppianthus microsporophyll sites, only the Lit 111
Dic/Hei TC yields carbonised compression material with the potential for
extracting in situ pollen. And only two of the nine available strobili
(PRE/F/5939 & PRE/F/5734) from this site appear to bear a few micro-
sporangia that could possibly yield pollen.
Cuticles
Potential sample: Lit 111,9 indivs.
Macerated (this work): 2 indivs.
Preservation grade: Grade 3 (fair), a few features available, small pieces
mainly from stalk area.
Diagnostic characters: cells oblong to narrowly oblong, walls straight to
gently curved; stomata absent; trichome bases apparently indicated by
circular marks, distribution random but towards the end of cells, on
both surfaces.
Comment: —
Significance:
Classification— The cuticular features do not indicate any particular
plant group, but also do not dispute placement in the Ginkgoopsida and
family Kannaskoppiaceae.
Affiliations— The diagnostic stomata and characteristic (?)glands
found in the leaf ( Kannaskoppifolia ) cuticle have not been found in that of
the affiliated male strobilus. The cell outlines and walls, however, corres-
pond well with that characterising the foliage.
Adaptive radiation (Molteno diversity)
Evident diversity (four species) in the male fruit, Kannaskoppianthus,
is clearly less than in the foliage where 10 species are recognised. (The
female fruit is represented by only one species from one TC.) The diag-
nostic characters of the four male species lie in the size and forking of the
strobilus and in the number of microsporophyll pairs per limb.
The Molteno species, based on the following TCs/palaeodemes, are
mainly derived from Heidiphyllum thicket within Cycle 3.
K. lutinumerus—Lut 311 Hei elo (Lutherskop), 16 indivs
Heidiphyllum thicket; Cycle 3 (Mayaputi Member)
K. matatiparvus— Mat 111 Die dub (Matatiele). 3 indivs
Dieroidium riparian forest (immature); Cycle 2 (Indwe Member)
K. irregularis — Kan 112 Hei elo (Kannaskop), 5 indivs
Heidiphyllum thicket; Cycle 3 (Mayaputi Member)
K. telemagnus—Je\ 111 Hei elo (Telemachus Spruit), 2 indivs
Heidiphyllum thicket; Cycle 3 (Mayaputi Member)
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
PETRIELLALES
Kannaskoppianthus
292
G7‘ TRELITZIA 15 (2003)
Kannaskoppianthus lutinumerus J.M.And. & H.M.And.,
sp. nov.
Holotype
Specimen : PRE/F/11433a,b; pi. 110(3,7,9).
Assemblage (TC): Lut 311 Hei elo, Lutherskop.
Preservation : complete very clearly preserved strobilus, part and counterpart;
impression in thickly laminated, medium grey shale with moderate
cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens'. 16indivs.pl. 110(1-10).
Sister palaeodemes— 9 (best 6 listed)
Aas 1 1 1 Hei elo: 21 indivs (6 intact, 15 partial), highest yield (5 indivs/man-day).
Aas 411 Dic/Sph: 21 indivs (10 intact, 11 partial), microsporangia present.
Lit 111 Dic/Hei: 9 indivs (6 intact, 3 partial), cuticle, microsporangia present.
Aas 311 Hei elo: 4 indivs (1 intact, 3 partial), microsporangia present.
Tel 111 Hei elo: 2 indivs (1 intact, 1 isolated), microsporangia present.
Aas 211 Hei elo: 1 indiv. (intact), microsporangia present.
Specific diagnosis
A Kannaskoppianthus species with small once-forked strobili whose
limbs bear 8-10 microsporophylls per row.
Specific characters
Attachment', unknown.
Strobilus : small ( ca 16-18 mm long), regularly forked; microsporophylls
8-10 per row.
Microsporophyll'. narrowly concavely conate (ca 1 mm wide distally).
Etymology
lutinumerus— luti, for the type locality Lutherskop; numerus (Lat.), with
reference to the large number of microsporophylls per row.
Comment & comparison
This species is by far the most numerous and frequent of the four
recognised in our Molteno collections. It is closely similar in form to K.
matatiparvus but differs in its larger size and greater number of micro-
sporophyll units per strobilus limb.
Kannaskoppianthus matatiparvus J.M.And. & J.M.And.,
sp. nov.
Holotype
Specimen. PRE/F/3205; pi. 114(10).
Assemblage (TC): Mat 1 1 1 Die dub, Matatiele.
Preservation : intact strobilus, without counterpart; impression in thickly
laminated, olive-grey shale with moderate cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens : 3 indivs (2 intact, 1 partial), pi. 114(8-10).
Sister palaeodemes— nil.
Specific diagnosis
A Kannaskoppianthus species with small once-forked strobili whose
limbs bear 5 or 6 microsporophylls per row.
Specific characters
Attachment', unknown.
Strobilus'. small (ca 8-10 mm long), regularly forked; microsporophylls
3-6 per row.
Microsporophyll'. narrowly concavely conate (ca 1 mm wide distally).
Etymology
matatiparvus— matati, for the type locality Matatiele; parvus (Lat.), small.
Comment & comparison
K. matatiparvus is particularly rare (3 indivs) and infrequent (1 TC)
and is differentiated from K. lutinumerus in the reduced number of micro-
sporophyll units per limb.
Kannaskoppianthus
PETRIELLALES
d^TRELITZIA 15 (2003)
293
Kannaskoppianthus irregularis J.M.And. & H.M.And.,
sp. nov.
Holotype
Specimen'. PRE/F/201 14a,b; pi. 117(1-6).
Assemblage (TC): Kan 112 Hei elo, Kannaskop.
Preservation : intact strobilus attached to an axis, part and counterpart;
impression in very thin-bedded, moderately baked, medium grey shale
with poor cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 5 indivs (1 intact, 4 partial), pi. 117(1-6).
Sister palaeodemes— 1 (as listed)
Kom 1 1 1 Sph/Dic: 2 indivs (1 partial, 1 fragm.), both attached to shoot with
foliage), pis 108, 109.
Specific diagnosis
A Kannaskoppianthus species with medium-sized irregularly forked
strobili whose limbs bear 5 or 6 microsporophylls per row.
Specific characters
Attachment : strobili attached terminally to a stem (?long shoot).
Strobilus: medium (>26 mm long), irregularly forked; microsporophylls 5
or 6 per row.
Microsporophyll: broadly concavely conate (ca 2 mm wide distally).
Etymology
irregularis— referring to the irregular branching of the strobilus.
Comment & comparison
K. irregularis differs from the other Kannaskoppianthus species in the
irregularly branching strobili and in its mode of attachment. On the holo-
type, PRE/F/201 14a,b, the stem shows leaf scars but no intact leaves. The
specimen from Kom 111 (PRE/F/3231) consists of a stem with four
attached leaves and two strobili that appear to be attached just below the
bifurcation of their axes, i.e. showing no evidence of multiple branching (tf.
2). In size and shape this specimen is best placed in this species— but with
uncertainty. The length of the strobilus may be much shorter than in the
holotype, with only four pairs of microsporangia preserved as indicated in
the reconstruction (p. 290, tf. 5).
Kannaskoppianthus telemagnus J.M.And. & H.M.And.,
sp. nov.
Holotype
Specimen: PRE/F/7711; pi. 118(1,2,4—6).
Assemblage (TC): Tel 111 Hei elo, Telemachus Spruit.
Preservation: almost complete strobilus, without counterpart; impression
in thickly laminated, light olive-grey shale with poor cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 2 indivs ( 1 intact, 1 partial); pi. 118(1-6).
Sister palaeodemes— nil.
Specific diagnosis
A Kannaskoppianthus species with large strobili whose limbs bear 9 or
10 microsporophylls per row.
Specific characters
Attachment: unknown.
Strobilus: large (>40 mm long), fork unknown; microsporophylls 9 or 10
per row.
Microsporophyll: broadly convexly conate (ca 3 mm wide distally).
Etymology
telemagnus— with reference to the type locality Telemachus Spruit and to
the large size of this species.
Comment & comparison
K. telemagnus differs strongly from the other three Kannaskoppi-
anthus species in the far larger size of its strobili and microsporophylls. It
is affiliated with the distinctive large-leaved Kannaskoppifolia sp.H
(p. 297, tf. 2) which, with 30 individuals (from Tel 111), is quite common.
Two specimens of K. lutinumerus have also been collected from this same
Tel 111 TC.
PETRIELLALES
Kannaskoppianthus
294
C/TRELITZIA 15 (2003)
Kannaskoppifolia J.M.And. & H.M.And., gen. nov.
Type species
Kannaskoppifolia vincularis J.M.And. & H.M.And., sp. nov.
Kannaskop. Karoo Basin, S. Africa; Camian, Triassic.
Generic diagnosis
A ginkgoopsid leaf without distinct petiole and with cuneate to flabel-
late, entire to trifidly divided lamina with forking, subparallel anastomos-
ing venation.
Generic characters
Attachment', leaves (1 or 2) and strobili (2 or 3) borne irregularly on short
shoots.
Leaf, narrowly to broadly cuneate to flabellate; petiole not distinct; lami-
na entire to deeply segmented with few to numerous segments based
on a trifid rather than bifid plan; veins fine, forking, radiating, subpar-
allel and anastomosing to form a variously elongated mesh.
Cuticle', this vol., p. 296, tfs 1, 2.
Etymology
Kannaskoppifolia— referring to leaves from the Kannaskop locality.
Global range— many spp., Gondwana, Tr. (SCY-NOR).
First: Kannaskoppifolia ( Ginkgoites sp.), (Walkom 1925a); L. Newport Fm.,
Turrimetta Head, Sydney, Australia.
Last: Kannaskoppifolia ( Chiropteris copiapensis). (Solms-Laubach & Stein-
mann 1899); ?Fm., Quebr. La Temera, Copiapo, Chile.
Gondwana Triassic occurrence
Frequency (F): 23 degree squares (of the 84 across Gondwana).
Ubiquity (U); 3 continents (of 5 comprising Gondwana).
Diversity (D): 10 foliage species.
Abundance (A): <1% (the norm in Molteno TCs).
Longevity (L): 26 myrs (Spathian to Upper Norian).
Colonisation success: FUDAL rating 23/3/10/-/26 = 62.
Intermediate success (Grade 3); Kannaskoppifolia was the 5th most
prominent foliage genus in the Gondwana Triassic; it was ubiquitous,
diverse, long-lived and relatively frequent, but generally lacking in
abundance.
Endemism: certain species occur widely distributed on three continents,
while at the other end of the scale, three species from the Molteno are
single-assemblage endemics.
Molteno occurrence
Frequency (F); 25 TCs (of 100 sampled in the Molteno).
Diversity (D): 10 species.
Abundance (A): common (3-5%) in 2 TCs; occasional (2%) in 2 TCs; and
<1% in the other 21 TCs.
Habit: possibly herbaceous pioneers, from erect shrublets to climbers.
Preferred habitat: occupied a wide range of habitats, but principally
Dicroidium riparian forest, Heidiphyllum thicket and fern meadows of
riverine sandbanks and floodplain wetlands.
Affiliated organs
Female strobilus: Kannaskoppia— Grade 5 (Org. att.).
Male strobilus: Kannaskoppianthus — Grade 5 (Org. att.).
Classification & comparison
Suprageneric classification
As this is an attached leaf, the classification is based on the ovulate
strobilus Kannaskoppia (p. 286).
Intergeneric comparison
Such leaves were frequently classified as Chiropteris or Ginkgoites
prior to Retallack (1980a) placing them in Ginkgophytopsis. In view of the
proven affiliation of attached leaves with male and female strobili occur-
ring in the Molteno, we here erect the new genus Kannaskoppifolia. We
suggest that Ginkgophytopsis be restricted to Devonian and Carboniferous
leaves as dealt with by Hpeg (1967), who included the genus in the order
Palaeophyllales.
Other reticulate leaves from the Molteno, such as Gontriglossa. Gracili-
glossa and Cetiglossa. all have a distinct midrib similar to that in Glossop-
teris (Permian) and Sagenopteris (Jurassic) leaves.
Eoginkgoites Bock from the Upper Triassic Newark Group and the
Chinle Fm., USA (Ash 1976, 1977) has anastomosing venation and synde-
tocheilic stomata, but differs in the finer venation, a marginal vein, pinnate
form and its papillate cuticle. Ash (1976, p. 1329) suggests a relationship
with the Bennettitales rather than the Ginkgoales.
Gondwana Triassic (elaborated)
Kannaskoppifolia is a widespread element throughout the Gondwana
Triassic. It is recorded in the literature from eight ‘localities’ (ca 88 indivs
illustrated) in S. America (Chile, N. Argentina, S. Argentina) and 15 ‘local-
ities’ in Australasia (Queensland, NSW, Victoria, South Australia and New
Zealand). Many of the published illustrations are insufficiently dear to
show whether the venation anastomoses or not. In such instances one might
be confusing this genus with Sphenobaiera, which can appear superficial-
ly similar (especially where specimens tend towards a bifurcating rather
than trifurcating division of the lamina). The abundance of Kannaskoppi-
folia is rarely, if ever, clearly stated in the literature. The general impres-
sion is that it is uncommon, as in the Molteno.
Recent literature (adding to above text)
Since virtually completing our manuscript, two papers directly relevant
to Kannaskoppifolia have appeared: Herbst et al. (2001), Barone-Nugent et
al. (2003, in press).
Herbst et al. (2001) erect the new genus Rochipteris for ‘some
Gondwanic leaf species previously assigned to Chiropteris Kurr ex Bronn'.
They describe five species of Rochipteris, two being new, from the Upper
Triassic of Argentina and Chile. Kannaskoppifolia should likely become a
junior synonym of Rochipteris. However, the South American foliage
apparently yields no cuticle and there is no mention of female or male fruit
affiliates.
Barone-Nugent et al. (in press) adopt the name Rochipteris for two
new species and a third unnamed species from the Ipswich Coal Measures,
the Leigh Creek Coal Measures and the Springfield Basin of the Upper
Triassic of eastern Australia. As for the South American material, there oc-
curs no affiliated fruit such as described here from the Molteno, but cuticle
is illustrated and described for the Leigh Creek species, R. amplexicaulis.
This cuticle is, in many respects, very different from that which we find
characterising the Molteno species. In particular, the stomata appear not to
be transverse and the cells are isodiametric (almost square over the veins),
not oblong. Furthermore, there are six subsidiary cells and these are clear-
ly lappetate. The Molteno stomata are paracytic and nonlappetate. How-
ever, certain specialised cells (trichomes or glands) are remarkably similar.
The leaf morphology of R. amplexicaulis, with its markedly expanded
clasping base, is likewise rather different from the Molteno Kannaskoppi-
folia species. We suggest that this species, at least, may be better placed in
a new genus within the family Kannaskoppiaceae.
Kannaskoppifolia vincularis J.M.And. & H.M.And.,
sp. nov.
Holotype
Specimen: PRE/F/13521 ; pi. 106(1).
Assemblage (TC): Kan 111 Ast sp.A, Kannaskop.
Preservation: a portion of shoot with leaves attached, without counterpart;
impression, in thick-bedded, moderately baked, greenish grey silty
mudstone with very poor cleavage.
Reference palaeodeme
Assemblage (TC): Kan 1 1 1 Ast sp.A, Kannaskop.
Specimens: 5 sections of shoot with foliage and megasporophylls attached;
12 sections of shoot with foliage only; >25 detached leaves, generally
tom or twisted (pis 104, 106, 107).
Sister palaeodemes— nil.
Specific diagnosis
A Kannaskoppifolia species, found attached to a shoot, with entire to
partly segmented narrowly wedge-shaped lamina.
Specific characters
Attachment: leaves on long shoots (?new shoots) in irregular helical
arrangement; leaves on short shoots in fascicle with female strobili.
Leaf, narrowly wedge-shaped, of medium size (up to 70 X 20 mm); lamina
entire to partly divided into 3 or more segments; base sharply angled.
Etymology
vincularis— vinculum (Lat.), link, bond, with reference to the fruit and
foliage being found in organic connection at the type locality.
Comment & comparison
This species is similar in shape and size to leaves previously described
as Ginkgophytopsis lacerata by Retallack (1983), but the latter have not
been found attached.
Kannaskoppifolia
PETRIELLALES
TRELITZIA 15 (2003)
295
PETRIELLALES
Kannaskoppifolia
296
dA-RELITZIA 15 (2003)
Cuticles
Potential sample: Lit 111. 51 indivs; Umk 111, 42 indivs.
Macerated (this work): Lit 111, 35 indivs; Umk 111, 30 indivs; only a
few indivs yielded good cuticle.
Preservation grade: Grade 4, features clear, fair-sized pieces.
Diagnostic characters: cells isodiametric to oblong (linear over vein
areas), walls gently curved; stomata hypostomatic, nonpapillate,
interveinal, transversely orientated; subsidiary cells paracytic, non-
cutinised. nonlappetate, guard cells narrowly elliptic; specialised
cells isodiametric to circular, strongly cutinised.
Comment: The specialised cells may be trichome bases or, in appearing
strongly cutinised, glands. They occur scattered on the upper and
lower cuticle. Such cells have not previously been recorded from the
Molteno.
Significance
Classification— The transversely orientated, paracytic stomata make
the cuticle of Kannaskoppifolia quite unique amongst the Ginkgoopsida.
The only other taxon with clear paracytic stomata in the Molteno is
Gontriglossa verticillata placed in the Gnetopsida. The latter has digitate
amorphous cells and the stomata are orientated randomly in the inter-
veinal areas.
The different species of Kannaskoppifolia from Lit 111 and Umk
111 all exhibit distinctive cuticles. That from the single specimen of K.
vincularis from Lit 111 is used here to illustrate the generic characteris-
tics. The second species. K. sp.F from Lit 111, and K. sp.E from Umk 1 1 1
both show papillate epidermal cells. While the other common species
from Umk 111, AT. sp.C. has yielded only poorly preserved cuticle, it does
show the diagnostic stomata and specialised cells. The remaining two
species (each represented by one individual) yield potential cuticle but
have not been macerated or studied.
Affiliations— Cuticular correspondence between the leaf and male
strobilus is not clear on present evidence and affiliation is based on direct
attachment.
Adaptive radiation (Molteno species, see text on p. 23).
We currently recognise 10 species of Kannaskoppifolia (Tab. 58) as
illustrated here (only the type species is described). Very interestingly,
the species in many instances coincide with different habitats, suggesting
that they are biologically and ecologically distinct entities.
assemblages
(taphocoenosis)
Cal 211 Hei/Ast
Gre 121 Hei elo
111 Equ sp
Cyp 111 Die era
Boe 112 Die cor
Kan 112 Hei elo
" 111 Ast spA
Tel 111 Hei elo
Kom 111 Sph/Dic
Vin 111 Die odo
Lut 311 Hei elo
Kon 211 Ast 2spp
Pen 311 Hei elo
" 411 Hei elo
Kapil 1 Dic/Ris
Nuw 111 Dic^zub
Win 111 Hei elo
Hla 21 3 Die elo
Total JCs
Total individuals
•2
33
Umk 111 Die 2spp
42
- -! 19 -! 21 !
San 111 Die era
3
- -) - - 2
Matlll Diczub
2
_l _l _l _l _l
Lit 111 Dic/Hei
56
.1 -1 -i -1 -1
Aas 111 Hei elo
2
-l -i £.\
” 211 Hei elo
19
- is
" 311 Hei elo
26
- 25
” 411 Dic/Sph
150
-?50
29
66
-i 70
6
4|__4j_i; _5I14
5! 19| 19)177! %
1
1 7 ’
55
30
50
Tab. 58. Kannaskoppifolia, Molteno occurrence
Lit 111 PRE/F/5641
prep. no. 973
Kannaskoppifolia
K. sp.A
R3
all x2
Kannaskoppifolia
PETRIELLALES
TRELITZIA 15 (2003)
297
PETRIELLALES
Kannaskoppifolia
d^TRELITZIA 15 (2003)
(Kan 111 Ast spA)
Kannaskoppia vincularis
pi. 104
PETRIELLALES
d?TRELITZIA 15 (2003)
299
PETRIELLALES
pi. 105
Kannaskoppia vincularis
300
c/tRELITZIA 15 (2003)
Kannaskoppial Kannaskoppifolia
pi. 106
PETRIELLALES
301
d/TRELITZIA 15 (2003)
PRE/F/13536'x1
Kannaskop
(Kan 1 1 1 Ast spA)
mgM
PRE/F/13545
:no. uncertain Mm
l- -r- 7 '»*> .'tic S' *• •
pi. 107
Kannaskoppifolia vincularis
PETRIELLALES
302
d^TRELITZIA 15 (2003)
Kommandantskop
(Kom 1 1 1 Sph/Dic)
Kannaskoppianthus irregularis
pi. 108
PETRIELLALES
ctf TRELITZIA 15 (2003)
303
PETRIELLALES
pi. 109
Kannaskoppianthus irregularis
304
d?TRELITZIA 15 (2003)
Lutherskop
(Lut 311 Hei elo)
PRE/F/11429b
Holotype
PRE/F/1 1429b
Kannaskoppianthus lutinumerus
pi. 110
PETRIELLALES
(Sf TRELITZIA 15 (2003)
305
Lutherskop
(Lut 311 Hei elo)
PRE/F/11386a
PRE/F/11359
PRE/F/11384a
PRE/F/11384a
PRE/F/1 1358a
PETRIELLALES
pi. Ill
Kannaskoppifolia sp.D
306
d/TRELITZIA 15 (2003)
PRE/F/19541
Aasvoelberg
(Aas 111 Hei elo)
PRE/F/15203a
BP/2/4297a
PRE/F/ 19545
BP/2/4436a
PRE/F/15227
BP/2/4436a
PRE/F/22514
PRE/F/22514
(jxio
PRE/F/19547b 3
PRE/F/15227
Kannaskoppianthus lutinumerus
pi. 112
PETRIELLALES
307
d^TRELITZIA 15 (2003)
Kannaskoppianthus
lutinumerus
Aasvoelberg
(Aas 211 Hei elo)
PRE/F/15280a
tv r jmmt
PRE/F/15280b
PRE/F/1 5280ag|
PRE/F/15280b
Kannaskoppifolia sp.E
PRE/F/15278
PRE/F/15278
PETRIELLALES
pi. 113
Kannaskoppianthus! Kannaskoppifolia
308
d^TRELITZIA 15 (2003)
il PRE/F/9251a'
?SWIV -
; Jp
'k f
PRE/F/9251b
Kannaskoppianthus
matatiparvus
Matatiele
(Mat 1 1 1 Die dub)
PRE/F/12001a
PRE/F/12001b X40
PRE/F/3205
PRE/F/12001biPjX40
Holotype
Aasvoelberg
(Aas 311 Hei elo)
Kannaskoppifolia
sp.F
PRE/F/9083b
SI
Kannaskoppianthus
lutinumerus
PRE/F/12001b
PRE/F/12001a
-■ 1 Kannaskoppifolia
sp.E
PRE/F/19019
PRE/F/19407b
PRE/F/19075
Kannaskoppianthus! Kannaskoppifolia
pi. 114
PETRIELLALES
309
d^TRELITZIA 15 (2003)
I wM
1 i l
wm%
' i,/ A
r
PRE/F/12050a
i -. Jt
PRE/F/12050a
Aasvoelberg
(Aas 411 Dic/Sph)
PRE/F/21720
Kannaskoppianthus lutinumerus
■w
PRE/F/21717b p
PRE/F/12093
PRE/F/12092a
Kannaskoppifolia -M
PRE/F/20558a
PRE/F/12172a
PRE/F/12172b
pi. 115
Kannaskoppianthus! Kannaskoppifolia
PETRIELLALES
310
Q7 TRELITZIA 15 (2003)
4' <V.
PRE/F/18277b
PRE/F/18277a
Telemachus Spruit
(Tel 111 Hei elo)
PRE/F/18277b ;
Vill
PRE/F/18277a
PRE/F/18277b
PRE/F/18277a
p PRE/F/18277b
PRE/F/18277a
PRE/F/18277a
Kannaskoppianthus lutinumerus
pi. 116
PETRIELLALES
311
Kannaskop
(Kan 112 Hei elo)
PRE/F/201 14b
~.w
Holotype
PRE/F/20114a
PRE/F/201 14b fS.
.. ~ — — — • AsSH
PRE/F/201 14b
PRE/F/201 14a
PRE/F/201 14b
c/tREL1TZIA 15 (2003)
PETRIELLALES
pi. 117
Kannaskoppianthus irregularis
312
d/TRELITZIA 15 (2003)
Kannaskoppianthus telemagnus
pi. 118
PETRIELLALES
c/ TRELITZIA 15 (2003)
313
Telemachus Spruit
(Tel 1 1 1 Hei elo)
PETRIELLALES
pi. 119
Kannaskoppifolia sp.H
314
d/TRELITZIA 15 (2003)
GINKGOOPSIDA S.V.Meyen 1987
INCERTAE SEDIS order
INCERTAE SEDIS family
CetifrilCtUS J.M.And. & H.M.And., gen. nov.
Type species
Cetifructus bilateralis J.M.And. & H.M.And.. sp. nov.
Umkomaas Valley, Karoo Basin, S. Africa; Camian, Triassic.
Generic diagnosis
A putative ginkgoopsid with a linear planar axis bearing opposite to
subopposite pairs of simple sessile megasporophylls comprising a pack of
several tiny linear-lanceolate ovules.
Generic characters
Strobilus : simple, small (up to ca 30 mm long), linear, laminate; axis flat-
tened, with distinct midrib and flanges, strongly curved; megasporo-
phylls opposite to subopposite, sessile, borne singly and semi-erect on
truncate flattened flange projections.
Megasporophyll: apparently reduced to single small (ca 2 mm long), ellip-
tical structures; ovules/seeds tightly packed, several per unit.
Ovule/seed: tiny (ca 2 X 0.2 mm), linear-lanceolate, striate.
Etymology
Cetifructus— cetus (Lat.), whale, with reference to the type locality Umko-
maas (which means cow or whale in the local Zulu language); fructus
(Lat.), fruit.
Global range: 1 sp., Gondwana, Tr. (CRN).
First & last: Molteno Fm.
Gondwana Triassic occurrence
SAf— Karoo Basin, 1 TC.
Molteno occurrence
Frequency (F): 1 TC (of 100 sampled in the Molteno).
Diversity (D): 1 species.
Abundance (A): 2 indivs.
Umk 1 1 1 : 2 indivs in 400 man-hrs cleaving ( 1 per 20 man-days) extremely rare
With only two specimens from one TC, Cetifructus is one of the rarest
of all Molteno fruit taxa. It does, however, meet our minimum criteria for
inclusion as a newly named generic entity.
Affiliated organs
Male : unknown.
Foliage: unknown.
Classification & comparison
Suprageneric classification (family & order incertae sedis)
Cetifructus is unlike any other ovulate organ from the Molteno or
known to us elsewhere in the fossil record. As the megasporophylls are
possibly cupulate structures, we place the genus in the class Ginkgoopsida.
However, owing to the uncertainty of the morphology, it is left in order and
family incertae sedis.
Intergeneric comparison (Gondwana Triassic)
It is not impossible that Cetifructus is microsporangiate. The linear
ovules/seeds, as interpreted, could in fact be pollen sacs. If so, and if our
reconstructions (tfs 1, 2 adjacent) are correct, then there is a similarity
between this genus and Antevsia (pp. 154, 155).
Reconstructions
With only two specimens at hand, the R3 and R4 reconstructions are
tentative. Apart from the strobilus being planar with opposite to suboppo-
site megasporophylls— features that are clear in the proximal(?) third of the
holotype— the other illustrated and described characteristics are uncertain.
Strobilus
Though the simpler interpretation would be that the megasporophylls
are attached distally and horizontally on the short stout projections, there
are four pointers to the reconstruction preferred here: the apparent scars
towards the centre of the flattened projections; the suggestion of vascular
bundles leading to these scars but not beyond; the megasporangiate rem-
nants at one of the points of attachment seeming to radiate outwards from
a scar; and the megasporangia along one side of the holotype almost all
being folded back across the axis.
Megasporophyll( ? )
Whether these structures are compound as proposed— with a tight
cluster of linear seeds partially or fully detached— or simple with one ellip-
tical seed or cupule, is not certain. Since the material is extremely rare (1
specimen per 20 man-days cleaving), we have chosen not to macerate a
specimen in the hope of resolving the uncertainties.
Cetifructus
GINKGOOPSIDA/INCERTAE
c/trelitzia 15 (2003)
315
Cetifructus bilateralis j.M.And. & H.M.And., sp. nov.
Holotype
Specimen-. BP/2/1191; pi. 120(1-5).
Assemblage: Umkl 1 1 Die 2spp, Umkomaas Valley.
Preservation: incomplete strobilus without counterpart; compression in thinly
laminated, carbonaceous (good cuticle) moderately baked, dark grey
shale with good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 2 indivs (2 intact); pis 120(1-5), 121(1-5).
Sister palaeodemes— nil.
Specific diagnosis— as for genus.
Specific characters— as for genus.
Etymology
bilateralis— with reference to the two lateral rows of megasporangia.
Comment & comparison— see for genus.
Umk 111 BP/2/1189
Cuticles
Potential sample: Umk 111,2 indivs.
Macerated (this work): none.
Preservation grade: —
Diagnostic characters: —
Comment: In an attempt to obtain some information on the epidermal
structure of Cetifructus, the material was placed under a Jed Scanning
Microscope (5800 LV). Linear striations that are probably cell walls
could be seen in certain places (see tfs 3, 4 adjacent) but no stomata or
other structures were visible.
Significance: of no aid in classification and affiliation.
GONDWANA TRIASSIC, GEOSTRAT DISTRIBUTION
GINKGOOPSIDA/INCERTAE
Cetifructus
316
y^y TREL1TZ1A 1J (ZUUJ)
MW*
*g&V ■
ALL BP/2/1191
Holotype
Umkomaas Valley
(Umk 1 1 1 Die 2spp)
Cetifructus bilateralis
pi. 120
GINKGOOPSIDA/INCERTAE
317
C/TRELITZIA 15 (2003)
ALL BP/2/1189
Umkomaas Valley
(Umk 1 1 1 Die 2spp)
GINKGOOPSIDA/INCERTAE
pi. 121
Cetifructus bilateralis
318
d/TRELITZIA 15 (2003)
INCERTAE SEDIS class
ALEXIALES J.M.And. & H.M.And., ord. nov.
ALEXIACEAE J.M.And. & H.M.And., fam. nov.
Alexia J.M.And. & H.M.And., gen. nov.
Type species
Alexia urceolus J.M.And. & H.M.And., sp. nov.
Umkomaas Valley. Karoo Basin. S. Africa; Camian, Triassic.
Generic diagnosis
A putative gymnospermous ovulate strobilus of uncertain form, with
linear planer axes bearing opposite rows of many simple sessile megas-
porophylls consisting of spherical pitcher-shaped cupules.
Generic characters
Strobilus : simple, small (>60 X ca 5 mm), linear planar ‘catkin’, gradually
tapering; axis gracile, curved, free for >10 mm; megasporophylls
numerous, sessile, opposite to alternate, decreasing gradually in size
distally.
Megasporophyll. pitcher-shaped with funnel-like distal projection function-
ing as a stigmatic surface or micropyle; semiwoody rather than fleshy.
Ovule! seed : unknown; possibly single, completely enclosed and coinciding
closely in dimension to the megasporophyll.
Eponymy
Alexia— in honour of Alex du Toit, the great pioneering geologist of South
Africa and of continental drift, who made the first substantial collec-
tions from the Umkomaas locality (ca 1910-1917).
Global range: 1 sp., Gondwana, Tr. (CRN).
First & last: Molteno Fm.
Gondwana Triassic occurrence
SAf— Karoo Basin, 1 TC (6 indivs).
Molteno occurrence
Frequency (F): 1 TC (of 100 sampled in the Molteno).
Diversity (D): 1 species.
Abundance (A): 6 indivs (1 strobilus, 5 isolated ovules).
Umk 111: 6 indivs in 400 man-hours ( 1 per ca 1 man-days) extremely rare
Affiliated organs: unknown.
Classification & comparison
Suprageneric classification (Alexiaceae/Alexiales)
A possible consideration is that Alexia is a fern belonging to a new
advanced order with elaborately developed, pitcher-shaped indusia. The
strobilus would then be regarded as a modified/reduced fertile frond— with
the indusia attached directly to the midrib. However, the semiwoody aspect
(rather than fleshy) and the fact that cuticle was obtained (fern cuticle being
invariably delicate and difficult to isolate from Molteno fronds), point to
this strobilus belonging to the Pinophyta (gymnosperms). Like several
other new Molteno megasporangiate strobili. Alexia is unique and merits
being placed in a new family (Alexiaceae) and order (Alexiales) and pos-
sibly even a new class (not named here).
Intergeneric comparison (Gondwana Triassic)
It is quite uncertain whether this single specimen represents the entire
strobilus, or whether it is a detached megasporophyll. Our description
assumes the latter. In this case there are some overall architectural similar-
ities with Hlatimbia, which, however, we include in a separate unnamed
class of fern-like gymnosperms.
Reconstruction
Strobilus
Our reconstruction is based on the only strobilus available, specimen
PRE/F/18745a,b, pi. 122(1), which is ca 60 mm long and incomplete. As
the axis does not taper, its full length in life cannot be estimated. A distinct
petiole is indicated by the absence of cupules towards the broken base.
There is a slight decrease in size of the cupules distally. Isolated cupules are
considerably larger than those attached to the axis. This, perhaps, is an
indication of maturity or size range.
Cupules
The strobilus shows only one complete cupule, which is characterised
by a pronounced funnel-like projection, pi. 122(4, 7, 8), and a second
cupule with a partial projection. An isolated cupule shows a partially pre-
served funnel, pi. 123(1, 2). In the reconstruction (tf. 1 above) we show
most cupules with the funnel-like projection.
Comparisons beyond Gondwana Triassic
A remote resemblance may be found in Eophyllogonium , a seed-
bearing gigantopterid leaf from the Permian of China described by
Mei et al. (1992). Eophyllogonium has a distinct leaf lamina with
oval seeds attached along the margins. The single similarity is that
both bear possible pollen traps, but the conical projections of
Eophyllogonium are quite different from the funnel-shaped structures
of Alexia.
A further resemblance occurs with the pitcher-shaped cupules of
Schmeissneria microstachys (Kirchner & Van Konijnenburg-Van
Cittert 1994) from the Liassic of Germany. However, the cupules of
the Laurasian genus differ in being either sessile (if single) or peduncu-
late (when in pairs or more) and in bearing attached winged seeds.
These strobili have been found in attachment with linear leaves (cf.
Glossphyllum etc.) now also included in the genus Schmeissneria by
the same authors.
Alexia
ALEXIALES
Cv^TRELITZIA 15 (2003)
319
Alexia urceolus J.M.And. & H.M.And., sp. nov.
Holotype
Specimen-. PRE/F/18745a,b; pis 122(1-8), 123(7, 8).
Assemblage (TC): Umk 1 1 1 Die 2spp, Umkomaas Valley.
Preservation', large section of strobilus, part and counterpart; compression in
thinly laminated, carbonaceous (good cuticle), moderately baked, dark
grey shale with good cleavage.
Reference palaeodeme
Assemblage (TC): as for the holotype.
Specimens: 6 indivs (1 intact, 5 isolated), pis 122, 123.
Sister palaeodemes— nil.
Specific diagnosis— as for genus.
Specific characters— as for genus.
Etymology
urceolus (Lat.)— a small jug or pitcher, with reference to the shape of the cupu-
late megasporophylls.
Comment & comparison— see for genus.
prep. no. 1002x
pi 123(8)
A. urceolus
Umk 111 PRE/F/18745
Cuticles
Potential sample: Umk 111,6 indivs.
Macerated (this work): 1 indiv.
Preservation grade: Grade 3 (fair), cell outlines only, small pieces.
Diagnostic characters: cells oblong to linear, walls straight to gently curved
to sinuous; other features absent.
Comment: —
Significance:
Classification— No cuticular features that aid in the classification of Alexia
are available.
Affiliation— Remains unknown and the cuticle gives no further clues.
ct
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
INCERTAE/INCERTAE
Alexia
320
Q/T RELITZIA 15 (2003)
Umkomaas Valley
(Umk 1 1 1 Die 2spp)
PRE/F/18745a
Holotype
X10
Alexia urceolus
pi. 122
INCERTAE/INCERTAE
INCERTAE/INCERTAE
pi. 123
Alexia urceolus
322
Q7 TR E LI TZ I A 15 (2003)
INCERTAE SEDIS class
HLATIMBIALES J.M.And. & H.M.And., ord. nov.
HLATIMBIACEAE J.M.And. & H.M.And., fam. nov.
Hlatimbia J.M.And. & H.M.And., gen. nov.
Type species
Hlatimbia tommacleanii J.M.And. & H.M.And., sp. nov.
Hlatimbe Valley, Karoo Basin, S. Africa, Carnian, Triassic.
Generic diagnosis
A putative gymnospermous ovulate strobilus of planar pinnate form,
with linear lateral axes bearing opposite rows of many simple pedunculate
megasporophylls consisting of bivalved ovulate cupules.
Generic characters
Strobilus: compound, paniculate, relatively large (>200 X 65 mm), dorsiven-
tral; main axis gracile (2-3 mm diam.); megasporophylls numerous, in
opposite rows along linear planar secondary axis (ca 70 mm long),
with a foliar tip.
Megasporophyll : simple, pedunculate; ovuliferous cupule bivalved (1.8 X 1.7
mm), bilaterally symmetrical; valves unequal, palmate, shallowly
cupped, lower valve larger and more deeply and profusely lobed (up to
8 teeth) than the upper (up to 5 teeth).
Ovule: unknown.
Etymology
Hlatimbia— after the type locality Hlatimbe Valley.
Global range: 1 sp.. Gondwana, Tr. (CRN).
First & last: Molteno Fm.
Gondwana Triassic occurrence
SAf— Karoo Basin. 1 TC.
Molteno occurrence
Frequency (F): 1 TC (of 100 sampled in the Molteno).
Diversity (D): 1 species.
Abundance (A): 2 indivs.
Hla 213: 2 indivs in 60 man-hours (1 per 3 man-days) very rare
Affiliated organs
Foliage: Batiopteris pttlchella— Grade 2 (Mut. occ.).
Male strobilus: unknown.
A unique foliar tip is preserved on one of the lateral axes (tf. 2 adjacent)
of Hlatimbia. This shows a similar shape to Batiopteris leaves from the
same TC and hence our Grade 2 affiliation.
Classification & comparison
Suprageneric classification (Hlatimbiaceae/Hlatimbiales)
At first glance, Hlatimbia gives the impression of being a rather
bizarre form of fertile fern frond, but the bivalved cupules and the absence
of sori place the genus in the Pinophyta (gymnosperms). We regard
Hlatimbia as most likely an ovulate structure — neither ovules or pollen are
known— and place it in the class incertae sedis. The unique bivalved
cupules and general architecture of the strobilus merit placing it in the new
order Hlatimbiales and family Hlatimbiaceae.
Intergeneric comparison (Gondwana Triassic)
The nearest Molteno ovulate genus to Hlatimbia appears to be Alexia,
which, however, is sufficiently remote as to suggest a separate unnamed
class of fern-like gymnosperms.
Hlatimbia
HLATIMBIALES
d^TRELITZIA 15 (2003)
323
Hlatimbia tommacleanii j.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/8951a,b,c; pis 124(1-7), 125(7, 8).
Assemblage (TC): Hla 213 Die elo, Hlatimbe Valley.
Preservation : large portion of an incomplete strobilus, part and counter-
part; compression in thinly laminated, carbonaceous (poor cuticle),
medium dark grey shale with good cleavage.
Reference palaeodeme
Assemblage: as for holotype.
Specimens: 2 individuals; both fairly substantial portions of strobilus.
Sister palaeodemes— nil.
Specific diagnosis— as for genus.
Specific characters— as for genus.
Eponymy
tommacleanii— in memory of Tom Maclean, on whose farm the Hla 213
site occurs.
Classification & comparison— see for genus.
Reconstruction
Strobilus
This remarkable paniculate strobilus is shown in the reconstruction
opposite, tf. 1. The main axis in PRE/F/8951a (tf. 1 adjacent) shows four
lateral axes attached and further broken fragments to the upper left, possi-
bly indicating another two detached lateral axes. The R4 reconstruction
shows a basally and apically incomplete main axis bearing six lateral axes.
In the second specimen, PRE/F/1357, one lateral axis is clearly
attached and fragments of a further six detached lateral axes are also pre-
sent. The two specimens possibly belong to the same strobilus, but owing
to missing matrix, the fossiliferous slabs do not fit together. Should the two
specimens belong together, then the strobilus would have at least 11 lateral
axes and be ca 250 mm long.
Foliar tip
The curious foliar tip is seen only on one lateral axis on specimen
PRE/F/8951a, pis 124(1-3), 125(7, 8). We assume that in life the foliar tip
would occur on all lateral axes, as in the reconstruction.
Megasporophyll/cupule
The bivalved cupule structure has been determined by excavation into
the matrix and observing that the valves lie at two levels. Usually only one
side is readily visible, but many of the cupules are variously flattened and
show portions of both upper and lower valves, pi. 124(3, 4). The distal
lobes vary from the usual simple projections, pi. 124(5-7), to the more rare
bilobed projections, pi. 125(9).
Classification beyond Gondwana Triassic
Laurasia Triassic
There is some similarity to the male strobilus Ixostrobus
described by Schweitzer (1977) from the Alborz Mountains, Rhaetic
of Iran (see also Harris & Miller 1974). Ixostrobus has four pollen
sacs which are united to form a woody cup-shaped synangium. It is
not impossible that Hlatimbia could be a male strobilus. Even so, it
is still very different from Ixostrobus which has a spiral arrangement
of synangia along the axis.
Other ages
The bivalved cupules of Hlatimbia show some resemblance to
Leptostrobus from the Yorkshire Jurassic of England (see L. cancer,
Harris & Miller 1974). The latter genus has cupules with an upper
and lower valve, but it differs in all other respects, such as the
attachment to the axis, the presence of seeds and the smooth margin
of the cupule.
Both Ixostrobus and Leptostrobus have clear affiliation to
Czekanowskia leaves (unknown from the Molteno) and are placed in
the order Leptostrobales.
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
HLATIMBIALES
Hlatimbia
324
d/TRELITZIA 15 (2003)
Bdtiopteris J.M.And. & H.M.And., gen. nov.
Type species
Batiopteris pulchella J.M.And. & H.M.And., sp. nov.
Hlatimbe Valley, Karoo Basin, S. Africa; Camian, Triassic.
Generic diagnosis
A gymnospermous leaf of uncertain class with long gracile petiole and
fan-shaped bifidly dividing lamina with frequently anastomosing venation.
Generic characters
Attachment: unknown.
Leaf, fan-shaped, broadly cuneate to conspicuously auriculate; lamina entire
to deeply and serially bifidly divided; petiole distinct, long, gracile;
venation frequently anastomosing, forming an open mesh.
Cuticle: this vol., see opposite.
Etymology
Batiopteris— batia (Gr.), bush, with reference to Hlatimbe based on the Zulu
word for bush; pteris (Gr.), fern.
Global range: 7 spp., Gondwana, Tr. (LAD-CRN).
First: Batiopteris ( Chiropteris barrealensis ) (Frenguelli 1942); Barreal Fm.,
Quebrada de la Cortaderita, Barreal, N. Argentina.
Last: Batiopteris zeilleri, Molteno Fm.
Gondwana Triassic occurrence
Frequency (F): 7 degree squares (of the 84 across Gondwana).
Ubiquity (U): 3 continents (of 5 comprising Gondwana).
Diversity (D): 7 species.
Abundance (A): <1% (as recorded for the Molteno).
Longevity (L): 2 myrs (Lower Camian).
Colonisation success: FUDAL rating 7/3/7/— /2 = 1 9.
Limited success (Grade 2); Batiopteris was the 18th most prominent
genus in the Gondwana Triassic; it was relatively ubiquitous and
diverse, but was infrequent, everywhere rare, and of apparently very
short longevity.
Endemism: In the thinly scattered nature of the palaeodemes and the rarity
of specimens, endemism was characteristically high in Batiopteris.
Four of the five Molteno species are single-assemblage endemics.
Molteno occurrence
Frequency (F): 10 TCs (of 100 sampled in the Molteno).
Diversity (D): 5 species.
Abundance (A): occasional (2%) in 1 TC; vanishingly rare (<1%) in 9 TCs.
Habit: possibly a slender, twining, herbaceous climber.
Preferred habitat: each species occurred in a different habitat, from ripar-
ian forest to Heidiphyllum thicket and fem meadow.
Affiliated organs
Female strobilus: Hlatimbia— Grade 2 (Mor. occ.).
Male strobilus: unknown.
Classification & comparison
Suprageneric classification
Batiopteris pulchella , and its affiliated megasporophyll Hlatimbia , are
tentatively placed in a new order and family.
Intergeneric comparison
Chiropteris, originally described from the Triassic in Germany, is a hetero-
genous group of leaves (Retallack 1980a) with anastomosing venation and
similar leaf shape to Batiopteris. However. Batiopteris is regarded as distinct
in view of affiliation with Hlatimbia and occurrence in Gondwana.
Gontriglossa, Cetiglossa , Gracilliglossa and Kannaskoppifolia all have
anastomosing venation but differ from Batiopteris in the form of venation
and lamina shape. Some species of Ginkgoites are similar to Batiopteris in
leaf shape but differ by the lack of anastomoses in the venation.
Additional Molteno leaves here listed as Batiopteris sp.A to sp.C will
be formally described in a future publication.
Gondwana Triassic occurrence (elaborated)
Batiopteris occurs very infrequently and rarely in South Africa and
elsewhere in Gondwana Triassic floras. Many leaves previously placed
in Chiropteris can now be included in Kannaskoppifolia or Rochipteris
Herbst et al. 2001. Those now regarded as Batiopteris are:
South America ( N . Argentina)
1942 Frenguelli: Chiropteris barrealensis , tf. 1, 2, pi. 1, 2(1, 2); a species
based on four illustrated specimens; one ‘locality’ (Quebrada de la Corta-
derita), Barreal-Hillario Basin, Barreal Fm., L. Camian, U. Triassic.
1963 Bonetti (unpubl. thesis): Chiropteris barrealensis, pis 8(1, 2), 10(1, 2),
32(6-8); seven illustrated individuals; one ‘locality’ (Punto 1 1 ), Barreal-
Hillaria Basin, Barreal Fm.
Australia ( Tasmania )
1888 Johnston: Sagenopteris salisburioides, f. 28 (4, 4A); a species based
on two very poorly illustrated individuals; one locality (Lords Hill),
Hobart, Brady Fm. equivalent, Camian, U. Triassic.
South Africa ( Karoo Basin )
1903 Seward: Chiropteris zeilleri, tf. 7; a species based on a single specimen
from Cyphergat, Molteno Fm. (illustrated here, tf. 5, opposite).
Batiopteris pulchella J.M.And. & H.M.And., sp. nov.
Holotype
Specimen' PRE/F/8697a,b; pis 126(16, 29), 127(3).
Assemblage (TC): Hla 213 Die elo, Hlatimbe Valley.
Preservation: complete leaf, part and counterpart; compression in thinly
laminated, carbonaceous (poor cuticle) medium dark grey shale with
good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 43 indivs.
Sister palaeodemes— nil.
Specific diagnosis
A Batiopteris leaf of small size with distinct petiole, moderately auricu-
late proximal lamina margin and variously divided distal lamina margin.
Specific characters
Leaf, small, ca 0.5-20 mm long and 0.5-30 mm wide; petiole distinct, ca
0.6 mm long and 0. 5-1.0 mm wide; lamina variously divided (in
smaller leaves once, in larger leaves up to 4 times), lobes obtuse; veins
in larger leaves ca 1 mm apart, with forks and anastomoses at ca
0.5-0. 8 mm intervals.
Etymology
pulchella (Lat.) — beautiful, with reference to the attractive leaves.
Comment & comparison
This species differs from other Batiopteris species in the combination
of size, shape and venation.
assemblages
(taphocoenosis)
Batiopteris j
B. pulchella (Hla 213)
” zeilleri (Cyp 111)
" sp.A (Win 111)
" sp.B (Kon 221)
” sp.C (Tel 111)
" spp. indet.
O Hlatimbia
Bir 111
Sph 2spp
1
1(11
-1 -i -1 -1 1
-
Cyp 111
Die era
20
- :20 1 - ! - ! - ! -
-
Tel 111
Hei elo
1
- ! -I
-
Kon 223
Die odo
1
-i - ; - : - ; -; i
-
Kon 221
Ast 2spp
2
-! -1 - ! 2 i - 1 -
-
Kon 211
Ast 2spp
6
- \ - 1 -i 6 - 1 -
-
Win 111
Hei elo
1
- ! - ! 1 1 - 1 - 1 -
-
Hla 213
Die elo
2
2 . 1 . 1 - 1 . 1 .
2
Umk 111
Die 2spp
2
-i - 1 - 1 - | - ; 2
-
Aas 311
Hei elo
1
- ! - > - 1 - 1 -i i
-
Total TCs
10
i i i i i i 2 i i i 4
1
Total indivs
%
% 20 1 8 1 5
2
Tab. 59. Batiopteris, Molteno occurrence
Batiopteris
HLATIMBIALES
G7TRELITZIA 15 (2003)
325
Cuticles
Potential sample: Umk 111,2 indivs; Hla 213, 43 indivs.
Macerated (this work): Umk 111, 2 indivs.
Preservation grade: Grade 2.
Diagnostic characters: cell structure present, but without diagnostic fea-
tures. ^
Comment: minimal interpretable structure obtained. <
Significance: of no value in classification or affiliation. £
GONDWANA TRIASSIC, GEOSTRAT’ DISTRIBUTION
HLATIMBIALES
Batiopteris
326
t v’; 1
PRE/F/8951a
PRE/F/8951a
PRE/F/8951a
Flolotype
Hlatimbe Valley
(Hla 213 Die elo)
PRE/F/8951a
H la ti in bia tonunacleunii
pi. 124
HLATIMBIALES
PRE/F/8951a
327
c/tRELITZIA 15 (2003)
i n
PRE/F/1357b
PRE/F/1357b
PRE/F/1357b
Hlatimbe Valley
(Hla 213 Die elo)
PRE/F/1357b
PRE/F/1357b
'■ PRE/F/8951a %
PRE/F/8951a
PRE/F/1357b
PRE/F/1357b
HLATIMBIALES
pi. 125
Hlatimbia tommacleanii
328
rV TRELITZI A 15
(2003)
8694
Holotype
8701
ALL PRE/F/ x2
Hlatimbe Valley
(Hla 213 Die elo)
8700
1182
8694
¥•
1191
8696
1179a'x'
\ 3
Holotype
1179b'y'
1179a'y'
8703
ALL PRE/F/
Batiopteris pulchella
pi. 126
HLATIMBIALES
329
HLATIMBIALES
pi. 127
Batiopteris pulchella
c/tRELITZIA 15 (2003)
PRE/F/8700
Hlatimbe Valley
(Hla 213 Die elo)
PRE/F/8701
PRE/F/8699
330
TRELITZIA 15 (2003)
INCERTAE SEDIS class
INCERTAE SEDIS order
INCERTAE SEDIS family
Hystncia J.M.And. & H.M.And., gen. nov.
Type species
Hystricia perplexa J.M.And. & H.M.And., sp. nov.
Aasvoelberg, Karoo Basin, S. Africa; Camian, Triassic.
Generic diagnosis
A gymnospermous ovulate structure of (?)dorsiventral circular form,
with a central 'gynoecium' of small ovuliferous cells surrounded by a perianth
of bracts.
Generic characters
Strobilus: compact, circular, bilaterally symmetrical, dorsiventral, medium
( ca 30 mm diam.); axis erect ( ca 1.5 mm diam.); fertile head consisting
of a central ‘gynoecium' fringed by a perianth of bracts.
‘ Gynoecium a honeycomb pack of numerous ovuliferous cells; bracts numer-
ous, irregular, leafy, free and overlapping to base, each extending from
a gynoecial cell.
Ovuliferous cell: minute (ca 0.5 mm diam.), pentagonal to hexagonal in surface
section, with a central depression or micropyle.
Ovule: unknown.
Etymology
Hystricia — hystrix (Gr.). porcupine, with reference to the spine-like bracts
surrounding the fruiting structure.
Global range: 1 sp., Gondwana, Tr. (CRN).
First & last: Molteno Fm.
Gondwana Triassic occurrence
SAf— Karoo Basin, 1 TC (1 indiv.).
Molteno occurrence
Frequency (F): 1 TC (of 100 sampled in Molteno).
Diversity (D): 1 species.
Abundance (A); 1 indiv.
Aas 411 Dic/Sph: 1 indiv. in 512 man-hrs (1 per 51 man-days) vanishingly rare
Though we have devoted 512 man-hours to cleaving slabs from this
highly significant lake-deposit site, no further sign of this uncertain taxon
has been found.
Hystricia perplexa J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/12936; pi. 128(1-5).
Assemblage (TC): Aas 41 1 Dic/Sph; Aasvoelberg.
Preservation: partial ‘gynoecium’, no counterpart; impression in thinly
laminated, strongly baked, yellowish grey shale with very good cleav-
age.
Reference palaeodeme— as for holotype.
Sister palaeodemes— nil.
Affiliated organs— unknown.
Species diagnosis— as for genus.
Classification & comparison
Suprageneric classification (incertae sedis/incertae sedis)
Hystricia, being represented by a solitary, fragmentary, unclear
impression fossil, can be classified only with considerable doubt. It bears a
superficial resemblance to the genus Ottokaria of the Permian
Ottokariopsida. It differs in having a fringe of large, entirely free bracts and
a head of numerous far smaller ovuliferous ‘cells’. There is also a resem-
blance to the Triassic Bennettitalean genera 'Williamsonia’ , Sturianthus
and Bennetticarpus, but these are all radially symmetrical and show a
peltate attachment of the pedicel. Although the features of Hystricia sug-
gest that it could be placed in the Ottokariopsida or the Bennettitopsida, we
place it in the class incertae sedis until more details are forthcoming.
Species description— as for genus.
Etymology
perplexa (Lat.)— confused, intricate, with reference to the enigmatic struc-
ture of the only available specimen.
Comment & comparison— see for genus.
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
Hystricia
INCERTAE/INCERTAE
331
Q/TRELITZIA 15 (2003)
INCERTAE/INCERTAE pi. 128 Hystricia perplexa
332
(ts TRELITZIA 15 (2003)
INCERTAE SEDIS class
INCERTAE SEDIS order
INCERTAE SEDIS family
Saportaea Fontaine & I. C. White 1880
Type species
Saportaea salisbitrioides Fontaine & I. C. White 1880
West Virginia, USA; Pennsylvanian. Carboniferous.
Generic concept
A gymnospermous leaf of uncertain class with a long gracile petiole
and lamina divided into 2 arcuate entire or deeply incised lobes with promi-
nent inner-marginal vein and subparallel forking secondary veins.
Generic characters (based only on the 2 Gondwana Triassic species)
Leaf, small to large, bipartite, with 2 primary arcuate lobes subtended by
a long slender petiole; lamina entire or deeply divided into dichotomis-
ing narrowly oblong to linear segments; venation consisting of a
prominent vein following the inner arcuate lohe margin and closely
spaced lateral veins, arching, subparallel, forking but not anastomos-
ing.
Cuticle : see And. & And. (1989, p. 530); this vol., tf. 5 opposite.
Eponymy
Saportaea— after G. Saporta, a 19th century French palaeobotanist.
Global range: several spp., Pangaea, L.P-U.Tr.
Gondwana Triassic occurrence
Frequency (F): 3 degree squares (of the 84 across Gondwana).
Ubiquity (U): 3 continents (of 5 comprising Gondwana).
Diversity (D): 2 foliage species.
Abundance (A): <1% (the norm in Molteno TCs).
Longevity (L): 9 myrs (Lower Anisian to Lower Camian).
Colonisation success: FUDAL rating 3/3/2/— /9 = 17.
Limited success (Grade 2); Saportaea was the 19th most prominent
genus in the Gondwana Triassic; it was relatively ubiquitous, diverse
and long-lived, but infrequent and rare.
Endemism: the 2 species are single-assemblage endemics.
Molteno occurrence
Frequency (F): 1 TC (of 100 sampled in the Molteno).
Diversity (D): 1 species.
Abundance (A): 1 indiv.; vanishingly rare.
Habit: most likely herbaceous undergrowth.
Preferred habitat: Dicroidium riparian forest.
Affiliated organs
Unknown.
Classification & comparison
Intergeneric comparisons
Gondwana Triassic gymnosperm genera— Saportaea, whose repro-
ductive organs remain unknown, could conceivably be an early member of
the Dipteridaceae. It was placed by Stipanicic & Bonetti (1965) in the
Ginkgoales in view of the obvious parallels in the dichotomising nature of
the lamina and venation. However, the prominent vein that follows the
inner margin of the arcuate lobes clearly distinguishes it from Ginkgo- like
leaves. We prefer to place it under incertae sedis (order level). The very
fragmentary cuticular remains available show gently curved cell walls, but
no further details to suggest relationships.
Saportaea, with its bipartite leaf with two primary arcuate lobes sub-
tended by a long slender petiole, is unlike any other gymnospermous leaf.
Other genera— The basic architecture of Saportaea is found also in the
fern family Dipteridaceae (U. Triassic to present). The fronds of the latter
differ in that they bear sori, have lamina segments that are often pinnate,
and generally show reticulate venation. The member of the family that
superficially looks most like Saportaea is the extant Dipteris conjugata,
but with its bifurcating lamina and reticulate venation (Andrews et al.
1970, f. 265) it is quite distinct.
Interspecific comparisons
Saportaea is a rare, nondiverse genus recognised from the L. Permian
to U. Triassic of the USA, China, Australia and S. Africa. Superficially, the
various species appear generically related, but this cannot be confirmed
owing to lack of cuticular information or fruiting bodies. S. magnifolia , the
single species described from the Molteno, is separated mainly on gross
morphological characters (And. & And. 1989, p. 530).
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
Saportaea
INCERTAE/INCERTAE
of' TRELITZIA 15 (2003)
333
S. dichotoma
schematic reconstructions
from And. &And., 1989
redrawn after photos in Stip. & Bon., 1965
S. magnifolia
tfs 5,6 from
And. & And., 1989
INCERTAE/INCERTAE
Saportaea
334
(37 T R E L ITZIA 15 (2003)
INCERTAE SEDIS class
INCERTAE SEDIS order
INCERTAE SEDIS family
Linguifolium E.Arber 1913
Type species
Linguifolium lillieanum E.Arber 1913.
Mt. Potts, New Zealand; Ladinian, Triassic
Generic concept
A gymnospermous leaf of uncertain class with linear-elliptic lamina,
entire margins and a midrib with steep forking secondary venation.
Generic characters
Leaf, small to medium, linear-elliptic to narrowly elliptic or oblanceolate;
apex sharply acute to obtuse; lamina entire, tapering very gradually to
base; midrib strong, tapering out short of apex; veins moderately to
well spaced, at steep angle to midrib and curving slightly, forking once
or twice but not anastomosing.
Cuticle : see And. & And. (1989, p. 520); this vol., tfs 1,2 opposite.
Etymology
Linguifolium— lingua (Lat.), tongue; folium (Lat.), leaf.
Global range: 5 spp., Gondwana, L.-U. Tr. (SCY-NOR).
First: Linguifolium sp. (Taeniopteris) (Walkom 1925a); Turrimetta Head,
L. Newport Fm., Sydney, Australia.
Last: Linguifolium arctum (Retallack 1985); Highfield Homestead, ?Fm.,
Nelson Syncline, New Zealand.
Gondwana Triassic occurrence
Frequency (F): 18 degree squares (of the 84 across Gondwana).
Ubiquity (U): 4 continents (of 5 comprising Gondwana).
Diversity (D): 5 foliage species.
Abundance (A): <1% (the norm in Molteno TCs).
Longevity (F): 25 myrs (upper Scythian to Upper Norian).
Colonisation success: FUDAL rating 18/4/5/-/25 = 54.
Intermediate success (Grade 3); Linguifolium was the 6th most promi-
nent genus in the Gondwana Triassic; it was of moderate frequency,
ubiquity and abundance, had marked longevity in the Triassic, but
lacked diversity.
Endemism: The five species may well come to be regarded as one wide-
spread polymorphic species in the future.
Molteno occurrence
Frequency (F); 8 TCs (of 100 sampled in the Molteno).
Diversity (D): 1 species.
Abundance (A): occasional (1%) in 1 TC; rare to extremely rare (<1%) in
other 7 TCs.
Habit: possibly a herbaceous pioneer.
Preferred habitat: Dicroidium riparian forest (5 of 8 TCs).
Affiliated organs (New Zealand)
Female strobilus: seeds only, Carpolithus mackayi , Grade 3 (Mut. occ.)
(see And. & And. 1989, p. 520).
Male strobilus: unknown.
Classification & comparison
Intergeneric comparisons
Gondwana Triassic genera— As a simple more or less linear to nar-
rowly elliptic leaf with clear midrib, Linguifolium is similar to Yabeiella
and Gontriglossa, but the former has simple bifurcating side veins while
the latter has anastomosing veins. The cuticle of Linguifolium , with straight
to gently curving cell walls, differs from Yabeiella and Gontriglossa which
both have meandering cell walls.
Other genera— Retallack (1980a) compared Linguifolium with a wide
range of somewhat similar genera of all ages and geographic origin.
Among those considered were Blechnoxylon Etheridge (wood with leaves
attached, ‘Permo-Carboniferous’, NSW, Australia), Phyllopteroides
Medwell (F. Jurassic, Victoria, Australia), Tatarina Meyen (U. Permian,
Kirovskaya Oblast, USSR), Lesleya Fesquereux (Pennsylvania, USA), and
Palaeovittaria Feistmantel (Permian, Raniganj, India). Retallack recog-
nised sufficient differences to treat Linguifolium as an independent genus.
Interspecific comparisons
The five species of Linguifolium recognised from the Gondwana
Triassic (And. & And. 1989) fall readily within the compass of a natural
genus on the basis of their leaf macromorphology. The available
palaeodemes of the five species present a complex series of overlapping
morphological ranges and undoubtedly represent a single genus if not a
single polymorphic species.
Cuticle is known only for the Molteno species L. gracile, while the
affiliated seed ( Carpolithus mackayi), apparently well established
(Retallack 1980a), remains known only for the New Zealand species.
GONDWANA TRIASSIC, GEOSTRAT’ DISTRIBUTION
Linguifolium
INCERTAE/INCERTAE
<§f'. TRELITZIA 15 (2003)
335
INCERTAE/INCERTAE
Linguifolium
336
^/tRELITZIA 15 (2003)
BENNETTITOPSIDA
FREDLINDIALES J.M.And. & H.M.And., ord. nov.
FREDLINDIACEAE J.M.And. & H.M.And.. fam. nov.
Fredlindia J.M.And. & H.M.And., gen. nov.
Type species
Fredlindia fontifructus J.M.And. & H.M.And., sp. nov.
Aasvoelberg, Karoo Basin, S. Africa; Camian. Triassic.
Generic diagnosis
A bennettitopsid ovulate strobilus of compact cone-like form; with
bilaterally symmetrical ‘gynoecia’ borne in a series of whorls.
Generic characters
Strobilus'. simple, compact, cone-like, medium (120 X 35 mm); axis stout
(7 mm diam. at base), markedly tapering; ‘gynoecia’ in ca 6 whorls of
3-8 units along axis.
' Gynoecium bilaterally symmetrical, tongue-shaped lamina, apparently
succulent, medium (25 X 10 mm); peduncle short, stout; ovuliferous
‘cells’ (megasporophylls) abaxial, a honeycomb aggregate of numer-
ous segments.
Ovuliferous ‘cell’: columnar, tapering gradually proximally, pentagonal to
hexagonal in section, (?)uni-ovulate; micropyle distinct, circular, central
with radial striae at distal face.
Etymology
Fredlindia— in honour of Fred and Linda Terblanche, on whose farm Aas
411 occurs.
Global range: 1 sp., Gondwana, Tr. (CRN).
First : Fredlindia sp. indet. ( Equisetites ? sp.) (Jones & De Jersey 1947); Tivoli
stage, Ipswich CM, Australia.
Last: Molteno Fm.
Gondwana Triassic occurrence
SAf— Karoo Basin, 3 TCs (16 indivs).
Aust— Clarence-Moreton Basin, 2 TCs (5 indivs).
Molteno occurrence
Frequency (F); 3 TCs (of 100 sampled in the Molteno).
Diversity (D): 1 species.
Abundance (A): 16 indivs total; very rare to extremely rare.
Kon 222 Die odo: 3 indivs in 40 man-hrs (1 per 1 man-day) very rare
Aas 411 Dic/Sph: 11 ” ” 512 ” (1 ” 5 ” ) extremely rare
Bir 111 Sph 2 spp: 2 ” ” 550 ” (1 ” 27 ” )
The potential for finding further Fredlindia specimens (other than at
new localities) rests largely at Aas 411. Although the yield there, to date,
has been very low— one specimen per five man-days cleaving— the site is
extensive, easy to excavate and the well-bedded sheets of hard shale are
readily cleaved. The yield at Bir 111, a very similar deposit, is five times
lower, while Kon 222, with a yield five times higher than Aas 411, is a site
of very limited extent and is apparently mined out.
Affiliated organs
Male strobilus : Cycadolepis/Weltrichia— Grade 3 (Mut. occ.).
Foliage : Flalleyoctenis— Grade 3 (Kin. reinf., Mut. occ.).
Classification & comparison
Suprageneric classification (Fredlindiaceae/Fredlindiales)
The Fredlindia ‘gynoecia’ are very like those of the Bennettitales in
being characterised by a honeycomb of ovuliferous ‘cells’ and/or intersem-
inal scales, but differ markedly in being bilaterally symmetrical and in
being borne in a series of whorls. Including the genus in the Bennettitales
would necessitate major alterations to that well-defined and understood
order. We prefer to erect a new order for the genus Fredlindia and to
include it with the Bennettitales in the class Bennettitopsida.
If the affiliation of the foliage Flalleyoctenis with Fredlindia proves
true, then this phylogenetic position is further substantiated. The early ben-
nettitalean leaf genera Laurozamites and Pterophyllum of the Laurasian
Late Triassic are clear intermediates between Flalleyoctenis and the typical
bennettitalean leaves of the Jurassic (see further detail under
Flalleyoctenis).
Intergeneric comparison (Gondwana Triassic)
Fredlindia is unique.
Fredlindia
FREDLINDIALES
G^/tRELITZIA 15 (2003)
337
Reconstructions
Strobilus
The reconstruction (tf. 1 opposite) is based primarily on PRE/F/ 12749
(the holotype from Aas 411, pi. 129) and PRE/F/2270 (from Kon 222, pi.
134)— the former consisting of the distal five whorls through to the clear-
ly preserved apex of the strobilus, and the latter of the proximal whorl,
including the complete stalk. The drawing is a combination of these two
specimens and results in a strobilus with four perfect whorls and two irreg-
ular whorls to the tip. The number of whorls that would comprise an aver-
age strobilus is not known and could well be greater than that shown in our
reconstruction.
‘Gynoecium '
The original thickness of the individual ‘gynoecium’ is uncertain.
Though all strobili are preserved three-dimensionally, the ‘gynoecia’ are
mostly preserved without significant dorsiventral thickness. The Kon 222
specimens (pi. 134) suggest that they were fleshy in life and were flattened
during fossilisation. The cross section of the ‘gynoecium’ (tf. 3 opposite) is
based on these Kon 222 specimens.
Ovuliferous ‘cell1
The ‘cells’ are most readily interpreted as elongate circumseminal
scales/sheaths, apparently fleshy, embedding a long tubular micropyle and
an ovule of unknown shape and size— reminiscent of the bennettitaleans.
They have been reconstructed to show the possible position of the micro-
pylar tube and ovule (tf. 5 opposite). Whether these ‘cells’ all had ovules or
whether some proportion of them were sterile is not known.
Gondwana Triassic occurrence (elaborated)
Australia
The individuals recorded for Australia in the hypodigm (Tab. 61) are
all described in the original literature (Shirley 1898; Jones & De Jersey 1947;
Hill et al. 1965) as being of sphenophyte origin, either as cones, cone-scars
or nodal diaphragms. Although the published illustrations are poor, we con-
sider it very likely that the three Australian specimens all represent de-
tached megasporophylls of Fredlindia strobili.
The material derives from two localities in the Clarence-Moreton
Basin, Queensland: Denmark Hill (Locality 39 or 39A of Jones & De
Jersey 1947), and portion 179 of parish Chuwar (Locality 6), from the
Blackstone and Tivoli Stages respectively. It is unclear from the descrip-
tions whether any of the specimens are compressions that might offer
potential for cuticular study.
Beyond Gondwana Triassic
The only bennettitopsid genus with ‘gynoecia’ attached to an
axis is Westerheimia from the Upper Triassic of Lunz, Austria. Crane
(1986) described W. pramelreuthensis as bearing ‘several bennetti-
talean “gynoecia”, each composed of interseminal scales and
ovules’. This differs from Fredlindia in lacking a whorled structure
and in its much smaller size.
Evidence for affiliation of organs
Mutual occurrence
Halleyoctenis is the most probable foliage affiliate of Fredlindia. It is
an abundant to extremely rare component of 10 Molteno TCs (Tab. 60).
Fredlindia is a very rare to extremely rare element, known from only three
Molteno TCs— each of which yields Halleyoctenis.
Kindred reinforcement
Supporting the evidence for ‘mutual occurrence’ is that both
Halleyoctenis and Fredlindia show certain features hinting clearly at a phy-
logenetic link with the order Bennettitales (see notes under ‘Classification’
opposite and on p. 344).
Australia
Halleyoctenis , as currently known, displays a markedly disjunct distri-
bution, appearing only in S. Africa and in Queensland. Fredlindia has the
same disjunct distribution, being unknown from any of the other
Gondwana continents.
The leaf genus Halleyoctenis is widespread in the Middle to Upper
Triassic formations of the Clarence-Moreton Basin, Queensland (And. &
And. 1989, p. 327, Tab. 4.38). It occurs in both the Denmark Hill and
Chuwar (Loc. 6) assemblages which yield the presumed Fredlindia speci-
mens (Jones & De Jersey 1947, pp. 45^48, 66, 70; And. & And. 1989, p.
365).
Intactness & preservation of cones (Molteno)
All but two of the 13 specimens are detached isolated megasporo-
phylls; PRE/F/2270 from Kon 222 consists of a proximal end of a strobilus
(stalk and lower whorl of megasporophylls); the holotype PRE/F/ 1 2749a,b
from Aas 41 1 consists of the greater part of a strobilus, but with the proxi-
mal end missing. The preservation in all three TCs is as 3D
impression/moulds .
Adaptive radiation (Molteno diversity)
The collections of Fredlindia, mostly detached ‘gynoecia’ from the
Molteno and two Queensland assemblages, are insufficient to justify the
recognition of more than one species. Considering the fact that three
species are currently recognised for the supposed foliage affiliate,
Halleyoctenis, it is likely that a more comprehensive sample of Fredlindia
material would reveal more diversity within the genus.
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
FREDLINDIALES
Fredlindia
338
^/treliTZIA 15 (2003)
Fredlindia fontifructus j.M.And. & H.M.And., sp. nov.
Holotype
Specimen-. PRE/F/12749a,b; pis 129(1-5), 130(1, 2).
Assemblage : Aas 411, Dic/sph, Aasvoelberg.
Preservation-, faidy complete strobilus, part and counterpart, including 5
whorls of ‘gynoecia'; proximal portion of strobilus not available (broken
off at a joint in the rock): impression in thinly laminated, strongly baked,
yellowish grey shale with very good cleavage.
Reference palaeodeme
Assemblage', as for holotype.
Specimens : 1 1 individuals; including the holotype and 10 isolated, detached
‘gynoecia’, pis 129-132.
Sister palaeodemes— 2 (both listed)
Kon 222 Die odo: 3 indivs (1 intact, 1 partial, 1 isolated), pi. 134.
Bir 111 Sph 2 spp: 2 indivs (2 isolated), pi. 133.
Specific diagnosis— as for genus.
Specific characters— as for genus.
Etymology
fontifructus— fontis (Lat.), source, spring, fountain; fructus (Lat.), fruit, with
reference to this fruit being near the fountainhead of the Bennettitop-
sida.
Comment & comparison
It is possible that the three available palaeodemes represent more than
a single species, but the material at hand is too sparse and incomplete to
make a judgement.
assemblages
(taphocoenosis)
Halleyoctenis {
H. brachypinnata
” megapinnata
.<TJ
*5
c
0)
u.
9
Q, Cycadolepis
Q, Weltrichia
Q, Leguminanthus
Bir 311 Hei/Sph
1
-i 1
-
” 111 Sph 2spp
6
3 3
2
1 < - ; -
Kon 223 Die odo
15
h-
00
-
" 222 Die odo
8
8 -
3
10 1 5
” 111 Die odo
10
10 -
-
- - ! -
Pen 321 Dic/Ris
5
5 -
-
1 _ l
” 211 Dic/Equ
2
2 -
-
- 1 - ] -
” 431 Dic/Equ
2
2 i -
-
- 1 - 1 -
Lit 111 Dic/Hei
8
-T 8
-
~1 2P
Aas 411 Dic/Sph
40
8 32
11
3 | - |
Total TCs
10
8 5
3
3 ! 2 ! 1
Total indivs
%
%1 %
16
14 ! 3 [ 5
Tab. 60. FredlindialHalleyoctenis,
Molteno occurrence
Tab. 61
FREDLINDIA HYPODIGM, Gondwana Triassic occurrence
Species
Intact-
ness
Mo.
F. fontifructus
<D
■o
C
d
a
w
u:
Intact strobili
Partial ”
Isolated megasp.
AUTHOR
SUBREGION
FORMATION
LOCALITY
NAME
Indivs ILLUSTRATION
AUSTRALASIA (Queensland, Clarence-Moreton Basin)
;
1 1
1 1
1 1
1898 1 Shirley
Ipswich/Esk CM5
24i Blackstone St
Denmark Hill
Equisetites mortonensis
1 f 18(3)
-
i
- i - ! i
1 947 Jones & de J.
’’ | ”
23! Tivoli St.
Chuwar (Loc.6)
Equisetites ?sp.
1 tf 1, pi 1(1)
-
i
- ! - 1 i
1965 Hill etal.
” | ”
24 Blackstone St
Denmark Hill
Neocalamites cf. c arrerei
1 plT1(3)
-
i
- ! - i
SOUTH AFRICA (Karoo Basin)!
1
l 1
1978-1999 Anderson & Anderson Molteno literature not included in this table
1
—
l 1
1
Fredlindia
FREDLINDIALES
c/tRELITZIA 15 (2003)
339
FREDLINDIALES
Fredlindia
340
d?TRELITZIA 15 (2003)
Weltrichia Braun 1847 (emend. T.M.Harris 1969)
Type species
Weltrichia mirabilis Braun 1847.
Origin of type material apparently obscure.
Generic characters
Flower’ (sensu Harris 1969, and others for ‘a Williamsonia fructifica-
tion’): cup-shaped with numerous (8-730) uniform tapering lobes.
Microsporophyll: an individual lobe; pollen sacs borne on inner surface.
Pollen sac: of 2 equal valves; microsporangia in a single row per valve,
opening inwardly.
Pollen: oval, monocolpate.
Etymology
Weltrichia— source of name unknown.
Global range: several spp., Pangaea, U. Tr.-U. K.
Gondwana Triassic occurrence
SAf— Karoo Basin.
Molteno occurrence
Frequency (F): 2 TCs (of 100 sampled in Molteno).
Diversity (D): 2 species.
Abundance (A): 3 indivs total; very rare to extremely rare.
Kon 222: 1 indiv. in 40 man-hrs ( 1 per 4 man-days) very rare
Lit 111: 2 ” ” 550 ” (1 ” 27 ” ) extremely rare
Affiliated organs
Female strobilus: Fredlindia— Grade 3 (Mut. occ.).
Foliage: Halleyoctenis— Grade 3 (Kin. reinf., Mut. occ.).
Classification & comparison
Suprageneric classification (Fredlindiaceae/Fredlindiales)
The Molteno material included here gives no direct indication of
whether this is a male or female structure. The specimens are placed in
Weltrichia (a male bennettitalean ‘flower’) on the basis of general similar-
ity. W. regalis is, perhaps, close to W. whitbiensis from the Yorkshire
Jurassic (Harris 1969, pi. 7, figs 4, 5).
Weltrichia regalis J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/3992a,b; pi. 135(1-3).
Assemblage (TC): Kon 222 Die odo, Konings Kroon.
Preservation: virtually complete ‘flower’, part and counterpart seen from outer
view, no details of inner pollen sacs visible; impression in thinly lam-
inated, medium grey cherty shale with poor cleavage.
Reference palaeodeme— 1 indiv. only (the holotype).
Specific diagnosis
A Weltrichia species with a broadly cup-shaped ‘flower’ (33 X 25 mm)
and ca 9 deeply lobed, lanceolate microsporophylls.
Specific characters
‘Flower’: broadly cup-shaped (33 X 25 mm), deeply lobed around distal peri-
meter into ca 9 microsporophylls.
Microsporophyll: lanceolate (to 20 X 5 mm).
Etymology
regalis (Lat.)— royal, with reference to the type locality Konings Kroon
(Afrikaans for King’s Crown).
Comment & comparison
The above description is based on the single available specimen. No
details of inner pollen sacs are visible.
Weltrichia helvetirara J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/22188; pi 135(4,5).
Assemblage (TC): Lit 111 Dic/Hei, Little Switzerland.
Preserx’ation: incomplete ‘flower’, base, tip and some of sides missing, no
counterpart, compression, some carbonised remains but mainly absent;
in thinly laminated, dark grey shale with moderate cleavage.
Reference palaeodeme— 1 indiv. only (the holotype).
Kon 222
PRE/F/3992b
pi 135(1-3)
Holotype
Lit 111
PRE/F/22188
pi. 135(4,5)
Holotype
W. helvetirara
2b,
possible
pollen sac
W. sp. indet
\vC^bT}//
Lit 111
BP/2/126P-
P' '35(6,7), , ^
xl
Specific diagnosis
A Weltrichia species with a narrowly cup-shaped ‘flower’ and at least
1 8 deeply lobed linear-lanceolate microsporophylls.
Diagnostic characters
‘Llower’: narrowly cup-shaped (726 X 16 mm), deeply lobed around distal
perimeter into at least 18 microsporophylls.
Microsporophyll: linear-lanceolate (up to 710 X 2 mm).
Etymology
helvetirara— helvetia (Lat.), Switzerland, with reference to the type locality;
rarus (Lat.), rare.
Comment & comparison
The description is based on the holotype alone. A possible pollen sac
(tf. 2b adjacent) occurs at the inner tip of a microsporophyll. but pollen has
not been extracted. A second specimen from Lit 1 1 1 [BP/2/1261 , tf. 3 adjacent,
pi. 1 35(6,7 )], differs in size and relief pattern and is probably a separate species.
GONDWANA TRIASSIC, GEOSTRAT DISTRIBUTION
SAm SAf Ind Ant Aus
Ch I NA I SA I Pa Lu I Za I Li Ka WH| PI A* NZ Ca SA Ga Bo CM| NE Sy Vi Ta
Weltrichia
FREDLINDIALES
^/tRELITZIA 15 (2003)
341
Cycadolepis Saporta 1873 (emend. T.M. Harris 1953)
Type species
Cycadolepis villosa Saporta 1873.
Orbagnoux, France; Triassic.
Generic characters
Dehisced bract : linear-lanceolate to arcuate, varying greatly in size (8-70
mm long), concavo-convex in transverse section, glabrous with reticu-
late wrinkled pattern or covered variously with hairs.
Etymology
Cycadolepis— lepis (Gr.), scale.
Global range: several spp., Pangaea, U. Tr.-U. K.
Gondwana Triassic occurrence
SAf— Karoo Basin.
Molteno occurrence
Frequency (F): 3 TCs (of 100 sampled in Molteno).
Diversity (D): 1 species.
Abundance (A): 14 indivs total; very rare to vanishingly rare.
Kon 222: 10 indivs in 40 man-hrs (2 per 1 man-day) very rare
Aas411: 3 " ”512 ” (1 ” 17 ” ) extremely rare
Bir 111: 1 ” ” 550 ” (1 ” 55 ” ) vanishingly rare
Affiliated organs
Female strobilus: Fredlindia— Grade 3 (Mut. occ.).
Foliage : Halleyoctenis— Grade 3 (Kin. reinf., Mut. occ.).
Classification & comparison
Cycadolepis is generally recognised as being an individual bract from
a caducous involucre enveloping hermaphroditic bennettitalean inflores-
cences (‘flowers’) such as Williamsoniella and Cycadeoidea (Menendez
1966; Harris 1969, p.102; Watson & Sincock 1992; Stewart & Rothwell
1993, pp. 352-361).
Cycadolepis rexiplumea J.M.And. & H.M.And., sp. nov.
Holotype
Specimen : PRE/F/20306a,b; pi. 136(4).
Assemblage (TC): Kon 222 Die odo, Konings Kroon.
Preservation', virtually complete scale, part and counterpart; impression in
thinly laminated, medium grey cherty shale with poor cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens'. 10 indivs (3 complete, 2 partial, 5 isolated), pi. 136(1-7).
Sister palaeodemes— 2 (as listed)
Aas 411: 3 indivs ( 1 complete, 2 partial ).
Bir 111: 1 indiv. (1 complete).
Specific diagnosis
A Cycadolepis species with linear-lanceolate scale (20->60 mm long),
bearing numerous long abaxial hairs.
Specific characters
Scale: linear-lanceolate, of variable size (20->60 mm long), sinuous to
curved or straight; hairs abaxial, dense, of a length approaching the
diameter of the scale, extending at acute angles.
Etymology
rexiplumea— rex (Lat.), king, with reference to the type locality; pluma
(Lat.), feather.
Comment & comparison
As no cuticles are known from Kon 222, close comparison with other
described species cannot be made. Based on occurrence and affiliation, the
Molteno material (from 3 TCs) is regarded as a new species.
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
FREDLINDIALES
Cycadolepis
342
^TRELITZIA 15 (2003)
Leguminanthus Krausel & F.Schaarschm. 1966
Type species
Leguminanthus siUquosus (Leuth.) Krausel & F.Schaarschm. 1966.
Neuevvelt. Basel, Switzerland; Keuper, U. Triassic.
Generic concept
A bennettitopsid microsporangiate organ consisting of a broad oblong
leaf-like pedunculate microsporophyll with laminae that are folded about a
midrib and bear microsporangia arranged in rows on the inner surface.
Generic characters (Molteno Fm.)
Microsporophyll : leaf-like, broad, oblong (45 x 8 mm); peduncle short and
broad; laminae folded about the broad midrib; venation parallel, lateral,
at a sharp angle to the midrib; microsporangia arranged in rows and/or
clusters on inner surface.
Microsporangium\ minute (<0.5 mm long), irregularly round to oval.
Etymology
Leguminanthus— legumen (Lat.), bean, with reference to the leguminous pod-
like appearance of the fruit.
Global range: ca 3 spp., Pangaea, U. Tr.
Gondwana Triassic occurrence
SAf— Karoo Basin.
Molteno occurrence
Frequency (F): 1 TC (of 100 sampled in Molteno).
Diversity (D): 1 species.
Abundance (A): 5 indivs; very rare.
Kon 222 Die odo: 5 indivs in 40 man-hrs (1 per 1 man-day), very rare
Affiliated organs
Female strobilus : unknown.
Foliage : see relevant discussion.
Classification & comparison
Suprageneric classification (Fredlindiaceae/Fredlindiales)
Based on northern Late Triassic collections, Leguminanthus is consider-
ed to be a bennettitalean pollen organ (Crane 1986, 1988). Although the
generic affiliation of this taxon within the Molteno is left open, it seems
most likely that it falls in the family Fredlindiaceae.
Intergeneric comparison (Gondwana Triassic)
Leguminanthus is unique.
Reconstructions
The reconstruction (tf. 3 opposite) is based on the holotype,
PRE/F/20410a,b, which shows good 3D preservation. Side ‘a’ clearly
shows the stalk, midrib area and the two distinct sides of the lamina; side ‘b\
the internal cast, shows the microsporangia more clearly, and in cross sec-
tion shows the incurving edge of the microsporophyll. In PRE/F/203 1 2b,
some of the microsporangia were prepared to reveal their spherical structure
(tfs la,b opposite). The venation, as seen on the type from Switzerland, is
not clearly preserved on the Molteno material and is not indicated in the
drawing.
Evidence for affiliation of organs
Bennettitopsid male organs occur extremely rarely in the Molteno
(Tab. 60). By far the most prolific TC, yielding all three genera recognised
( Cycadolepis , Weltrichia and Leguminanthus), is Kon 222. Cycadolepis
and Weltrichia are considered as affiliating with Fredlindia. The affiliation
of the Molteno Leguminanthus is uncertain at present.
We provide a comparative floristic synopsis from three localities
which suggests possible affiliation: the first, from the Molteno, is outlined
below; the other two, from the Late Triassic of Europe, are discussed in the
box at the foot of the page.
Kon 222 Die odo, Molteno Fm.
A typical TC representing the Dicroidium open woodland of the floodplain. It has
been well sampled, at 40 man-hours cleaving, and includes 449 catalogued slabs.
Bennettitales: This is the second most dominant gymnosperm element in the flora
after Dicroidium (at 87%).
Foliage— 1 genus. Halleyoctenis (8%).
Female fruit— the single new genus Fredlindia , with 3 individuals.
Male fruit— 3 genera of apparent bennettitopsid male organs occur; Cycadolepis ,
Leguminanthus and Weltrichia. Apart from this Kon 222 material, Cycadolepis
bracts are recorded from Aas 4 1 1 and Bir 111 and Weltrichia from Lit 111.
Various alternative interpretations concerning the nature and affilia-
tions of these three organ-genera exist:
a) All are elements of a single complex fruit affiliated to Fredlindia and
Halleyoctenis.
b) They represent three distinct organs of bennettitopsid plants, at least
two of which are without female or foliage affiliates at Kon 222.
c) Cycadolepis, a sterile bract, pairs with either Weltrichia (our prefer-
ence) or Leguminanthus, leaving at least one taxon without female or
foliage affiliates. [Note that the only other possible foliage affiliate is
Pseudoctenis, which is identified as cycad rather than bennettitopsid
on the basis of gross morphology and cuticle (And. & And. 1989, pp.
280-325).]
d) One or more of these supposed male elements is not bennettitopsid.
Comparison beyond Gondwana Triassic
Krausel & Schaarschmidt (1966) made a comparison of
Leguminanthus with other possible bennettitopsid pollen organs, e.g.
Williamsoniella. Leuthardtia, Hartingeria and Lunzia, and conclud-
ed that it was unique. Brightonia (Harris 1932b, p. 119, pi. 19), from
the Late Triassic Lepidopteris Zone of Greenland, was considered by
Crane (1986) to be similar to Leguminanthus.
On the bennettitopsid affiliation of Leguminanthus in the Late Triassic of Europe
Lutiz., 100 km .S'VL of Vienna, Austria (Carnian)
The plant beds, with several TCs over a couple of kilometres, occur
within a coal-mining area which is part of a limestone sequence in the
northern Alpine foothills. Marine invertebrates provide a Carnian age.
Apart from the general report of Dobruskina (1988), there is no thorough
documentation of abundance data per site for the Lunz collections. It is not
feasible, therefore, to reliably establish affiliations based on the co-occur-
rence of organs at TC level. However, the flora shows a high percentage of
Bennettitales (abundance and diversity), and Leguminanthus, with 22 indi-
viduals available, most likely affiliates with this order.
Foliage (based on the study by Dobruskina 1988):
The overall flora comprises sphenophytes (6%), ferns (7%), Bennettitales (ca
53%), Taeniopteris (17%), Macrotaeniopteris (<1%), ginkgophytes (13%) and
conifers (5%).
Bennettitales1. The dominant element of the flora, and the most diverse amongst
the gymnosperm orders.
Foliage— 4 species recorded: Pterophyllum longifolium (50%); Pterophyllum sp.B,
Anomozamites sp. and Nilssonia sturi (each > 1%).
Female fruit— 3 genera: Westerheimia, Sturianthus, Bennetticarpus.
Male fruit— 3 genera: Leguminanthus, Haitingeria, Cycadolepis.
Neuewelt, Basel, Switzerland ( Keuper )
Three localities, Neuewelt, Moderhalde and Hemmiken, occur
within or in the neighbourhood of Basel. Neuewelt, the best known and
described of the floras, comprises a number of TCs scattered around the
flanks of the Rutihard hill within the city. We confine our discussion to
this ‘locality’. There exists no synthesis of the flora providing abun-
dance data, so assessment of affiliations remains provisional. The link
between Leguminanthus and a species of Pterophyllum seems probable.
Foliage (based on unpubl. literature survey by H.M. Anderson, 1973):
The overall flora consists of two genera and five species of sphenophyte
(common to abundant), eight genera and 10 species of fem (rare to common),
Bennettitales (abundant), Taeniopteris, ginkgophytes (common) and conifers (?
rare).
Bennettitales: The dominant gymnospermous element in the flora.
Foliage— 1 genus: Pterophyllum (3 recorded species); the second most abundant
element in the flora after Equisetum.
Female fruit— unknown.
Male fruit— 2 genera: Leguminanthus (ca 3 indivs); Williamsonianthus keuperi-
antltus (based on a single specimen with 6 cupules).
Leguminanthus
FREDLINDIALES
^ TRELITZIA 15 (2003)
343
Leguminanthus leopardus j.M.And. & H.M.And., sp. nov.
Holotype
Specimen'. PRE/F/20410a,b; pis 137(3-5), 138(3-5).
Assemblage (TC): Kon 222 Die odo, Konings Kroon.
Preservation', a complete specimen, part and counterpart, longitudinal view;
3D mould and cast in thinly laminated, medium grey cherty shale with
poor cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 5 indivs (3 intact, 1 partial, 1 isolated), pis 137, 138.
Specific diagnosis
A Leguminanthus species with clusters of circular to oval microspor-
angia aligned in irregular diagonal to horizontal rows.
Specific characters (Molteno Fm.)— as for genus.
Etymology
leopardus (Lat.) — leopard, with reference to the appearance of the microspor-
angial groups (particularly in the unprepared specimens), reminiscent
of the markings of a leopard.
Comment & comparison
The Molteno specimens closely resemble the type species in general
morphology, but differ in the arrangement of the putative microsporangia
into irregular diagonally aligned clusters rather than in clear transverse
rows.
L. leopardus
microsporangia in distinct clusters
microsporangial clusters shown on inner surface
, , cross section
based on
PRE/F/20410a.b
Kon 222
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
FREDLINDIALES
Leguminanthus
344
d^TRELITZIA 15 (2003)
Halleyoctenis J.M.And. & H.M.And. 1989
Type species
Halleyoctenis multilineata (Shiriey 1897) J.M.And. & H.M.And. 1989.
Ipswich-Esk, Clarence-Moreton Basin. Queensland; Camian, Triassic.
Generic diagnosis
A bennettitopsid leaf with roundly truncate pinnae and fine, very closely
spaced, parallel, occasionally forking venation.
Generic characters (based on the three definite Gondwana species)
Leaf, small to large, obovate to narrowly elliptic, simply pinnate; pinnae
laterally to slightly dorsally attached, margins entire, oblong to nar-
rowly oblong, apex truncate, base slightly expanded or contracted;
veins numerous, very closely spaced, parallel, occasionally forking but
not anastomosing, terminating along the pinna margin and mostly
across apex.
Cuticle : see And, & And. (1989, p. 327); this vol., tfs 1-3 adjacent.
Eponymy
Halleyoctenis— after the English astronomer E. Halley, and the cycad leaf
Ctenis.
Global range: 3 spp., Gondwana, M.-U. Tr. (ANS-CRN).
First: Halleyoctenis sp. (Webb 1980); UQL.4224, Bryden Fm., Ipswich/Esk,
Australia.
Last: H. brachypinnata , Molteno Fm.
Gondwana Triassic occurrence
Frequency (F): 7 degree squares (of the 84 across Gondwana).
Ubiquity (U): 2 continents (of 5 comprising Gondwana).
Diversity (D): 3 foliage species.
Abundance (A); 2% (the norm as in Molteno TCs).
Longevity (L): 9 myrs (late Anisian to Lower Camian).
Colonisation success: FUDAL rating 7/2/3/2/9 = 24.
Limited success (Grade 2): Halleyoctenis was the 14th most prominent
foliage genus in the Gondwana Triassic; it enjoyed relatively low to
moderate frequency, ubiquity, diversity, abundance and longevity.
Endemism: of the 3 (+1; see Umk 111, BP/2/793, tfs 5, 6 opposite) described
Gondwana Triassic species, 2 are basin endemics and 2 are single-
assemblage endemics.
Molteno occurrence
Frequency (F): 10 TCs (of 100 sampled in the Molteno).
Diversity (D): 2 (+1; see Umk 111, BP/2/793, tfs 5, 6 opposite) species.
Abundance (A): abundant (8-15%) in 2 TCs; rare to extremely rare (<1%)
in 8 TCs.
Habit: probably cycad-like plants.
Preferred habitat: most common in Dicroidium open woodland, more rare in
closed woodland of the lake margin.
Affiliated organs
Female strobilus: Fredlindia— Grade 3 (Mut. occ.).
Male strobilus: CycadolepisIWeltrichia— Grade 3 (Mut. occ., Kin. reinf.).
Classification & comparison
Suprageneric classification
In many of its features, Halleyoctenis is the perfect proto-bennetti-
talean foliage. It has more in common with a number of the early members
of the Bennettitales as known from the Late Triassic of the northern hemi-
sphere than with the more typical forms of the Jurassic (e.g. Harris 1969).
Intergeneric comparison
Laurozamites Weber & Zamudio- Varela (1995) is the most common
element of the Late Triassic (Camian) Santa Clara Fm. of Sonora, Mexico.
It has been collected from many localities and several species are recog-
nised. Zamites powelli , now included in Laurozamites (Weber & Zamudio-
Varela 1995), is one of the most common leaves (from about 35 localities)
in the Camian to Lower Norian formations of the Late Triassic in the USA
(Ash 1975). Laurozamites is very like Halleyoctenis in the general appear-
ance of the frond, in the shape of the pinnae with their truncate apex, in the
very fine closely spaced, parallel to slightly spreading venation, and in sev-
eral features of the cuticle, such as the transverse stomata, the guard cells,
the shape of the cells both veinal and interveinal, and the single papilla per
cell. It is more bennettitalean than Halleyoctenis in the strongly contracted
base of the pinnae and their marked dorsal attachment, and in critically
diagnostic features of the cuticle such as the mildly sinuous to meandering
cell walls, the pair of well-differentiated subsidiary cells and the greater
stomatal frequency.
Certain Late Triassic species of the bennettitalean genus Pterophyllum
also have much in common with Halleyoctenis. Examples include P.
aequale from the Donetz Basin, Russia (Stanislavsky 1976), and P. spp.
from the Hunan Province, China (Zhou Zhiyan 1989). These bear oblong
pinnae with truncate apices and appear, at least partly, to be laterally
attached and are not contracted at the base. They differ from Halleyoctenis
in their more widely spaced veins. The cuticle of the Chinese species, with
oblong cells and transverse stomata, although with greater stomatal fre-
quency and mildly meandering cell walls, is similar to that of Laurozamites
and is transitional between Halleyoctenis and the Jurassic Bennettitales.
Based on both mega- and micromorphological characters, Halley-
octenis is readily differentiated from the four Gondwana Triassic cycad
genera. It is perhaps nearest to Pseudoctenis in frond morphology, but dif-
fers most notably in the transverse orientation of the stomata.
Interspecific comparison
Four species are recognised in the Gondwana Triassic (And. & And.
1989). The two Molteno species, H. brachypinnata and H. megapinnata,
and the Australian species H. multilineata are closely similar— their distin-
guishing features being the shape and size of the frond and the individual
pinnae. Cuticle is known only for H. megapinnata. The single specimen
from Umk 1 1 1 (BP/2/793, tfs 5, 6 opposite), described in our earlier work
as H. symmetrica , should very probably be transferred to a new genus.
The Molteno collections of Halleyoctenis have at least doubled (Tab.
60) in assemblages (five new TCs) and individuals since our original
description of the genus (And. & And. 1989), and an updated systematic
comparative study of the full set of palaeodemes is due.
Herbst & Troncoso (2000) describe two specimens as Pseudoctenis
multiline atum from locality 274, near Copiapo, La Temera Fm., Chile.
Their illustrated specimen showed a small frond fragment with venation
that we consider insufficiently preserved to verify their identification.
Herbst & Troncoso have not accepted our transfer (And. & And. 1989) of
Pterophyllum multilineata Shirley to the bennettitalean genus
Halleyoctenis, although that decision was based on good cuticular evidence
from the Molteno. Instead, they place their specimen, without preserved
cuticle in support, in the cycad genus Pseudoctenis.
Halleyoctenis
FREDLINDIALES
d/TRELITZIA 15 (2003)
345
tfs 1-6 from
And- & And . 1989
H. brachypinnata
Halleyoctenis
generic panorama
showing the 2 (+1 )
Molteno species
This species, included as Halleyoctenis in
And.& And., 1989, is here considered to very
propably represent a new unnamed genus
GONDWANA TRIASSIC. GEOSTRAT’ DISTRIBUTION
FREDLINDIALES
Halleyoctenis
346
TRELITZIA 15
(2003)
Holotype
Aasvoelberg
(Aas 411 Dic/Sph)
Fredlindia fontifructus
pi. 129
FREDLINDIALES
347
^/tRELITZIA 15 (2003)
FREDLINDIALES
pi. 130
Fredlindia fontifructus
348
G?^T RELITZIA 15 (2003)
Fredlindia fontifructus
pi. 131
FREDLINDIALES
d^TRELITZIA 15 (2003)
349
Aasvoelberg
(Aas 411 Dic/Sph)
PRE/F/2 1434a m
PRE/F/21434b
PRE/F/2 1434b
PRE/F/2 1434b
PRE/F/2 1434b
FREDLINDIALES
pi. 132
Fredlindia fontifructus
350
(§? X
RELITZIA 15 (2003)
Birds River
(Bir 1 1 1 Sph 2spp)
Fredlindia fontifructus
pi. 133
FREDLINDIALES
351
(§?'. TRELITZIA 15 (2003)
FREDLINDIALES pi. 134 F redlindia fontifructus
352
TRELITZI A 15 (2003)
Weltrichia spp. pi. 135 FREDLINDIALES
353
d/ TRELITZIA 15 (2003)
PRE/F/20308b ; X2
BP/2/4130
Konines Kroon
(Kon 222 Die odo)
PRE/F/20307a
^R^^20306b ^
Holotype
PRE/F/20307a
PRE/F/20307a
BP/2/4130
FREDLINDIALES
pi. 136
Cycadolepis rexiplumea
354 d/ TRELITZIA 15 (2003)
Konings Kroon
(Kon 222 Die odo)
Leguminanthus leopardus pi. 137 FREDLINDIALES
355
c/tRELITZIA 15 (2003)
pi. 138
Konings Kroon
(Kon 222 Die odo)
Holotype
xlO
FREDLINDIALES
Leguminanthus leopardus
356
& TRELITZIA 15 (2003)
BENNETTITOPSIDA
PENTOXYLALES Pilg. & Melch. 1954
LINDTHECACEAE J.M.And. & H.M.And., fam. nov.
Lindtheca J.M.And. & H.M.And., gen. nov.
Type species
Lindtheca hackysackia J.M.And. & H.M.And., sp. nov.
Aasvoelberg. Karoo Formation, S. Africa, Camian. Triassic.
Generic diagnosis
A pentoxylalean ovulate structure comprising a spherical ‘gynoecium’
with 15-20 ovuliferous 'cells’ (megasporophylls) each containing 5 ovules.
Generic characters
Strobilus : unknown
‘ Gynoecium radially symmetrical, spherical pedunculate balls, apparently
succulent, small (ca 10 mm diam.); with an aggregate of 15-20 pentago-
nal segments or ovuliferous cells (megasporophylls); an outer ‘skin’,
with well-defined puckered sutures that follow the pentagonal out-
lines of each segment, covers the ‘gynoecium’; peduncle known only
through clear attachment scar, evidently gracile.
Ovuliferous ‘cell’', prism-shaped, radiating from a central receptacle, irregu-
larly pentagonal in section; multi-ovulate, apparently with 5 seminal
scales per ‘cell’ each bearing an ovule; micropyle distinct, circular, off-
centre, with a clear pattern of radiating cells on outer skin.
Eponymy
Lindtheca— in honour of Linda Terblanche, on whose farm the type locality
Aasvoelberg (Aas 411) occurs.
Global range: 1 sp.. Gondwana, Tr. (CRN).
First & last. Molteno Fm.
Gondwana Triassic occurrence
SAf— Karoo Basin, 1 TC (16 indivs).
Molteno occurrence
Frequency (F): 1 TC (of 100 sampled in the Molteno).
Diversity (D): 1 species.
Abundance (A): 16 indivs; very rare.
Aas 411 Dic/Sph: 16 indivs in 512 man-hrs (1 per 4 man-days) very rare
Affiliated organs
Male strobilus'. unknown.
Foliage : Taeniopteris homerifolius— Grade 3 (Kin. reinf., Mut. occ.).
Classification & comparison
Suprageneric classification (Lindthecaceae/Pentoxylales)
The pentoxylalean ovulate genus Carnoconites (Crane 1985) is widely
known in the Jurassic and Lower Cretaceous of India (Bose et al. 1985),
Australia (White 1981; Drinnan & Chambers 1985) and New Zealand
(Harris 1962, 1982). Carnoconites is a unique fructification, which has
spherical ‘gynoecia’ with numerous uni-ovulate ‘cells’. Lindtheca is similar
to Carnoconites in all features with the exception that it is multi-ovulate.
On this basis we place Lindtheca in the order Pentoxylales and in the new
family Lindthecaceae. Lindtheca is the first genus allied to Carnoconites to
be described from the Triassic.
Intergeneric comparison (Gondwana Triassic)
Lindtheca is unique.
Evidence for affiliation of organs
Kindred reinforcement
Nipaniophyllum is a Taeniopteris-like leaf that has been affiliated with
cones and stems and placed in the order Pentoxylales (Sahni 1948; Stewart
& Rothwell 1993). Lindtheca , placed in the Pentoxylales, is thus more likely
to be affiliated with a Taeniopteris-like. leaf.
Mutual occurrence
The 16 individuals of Lindtheca— all isolated ‘gynoecia’— derive from
a single TC, Aas 411. Of all the gymnosperm foliage genera found at Aas
411, Taeniopteris is the most likely affiliate. All other gymnosperm leaf
genera have been assigned affiliates except for a Pseudoctenis species here
classified as a cycad. Cuticle, not preserved at Aas 411, is unavailable to
confirm affiliations.
Lindtheca
PENTOXYLALES
d/TRELITZIA 15 (2003)
357
Reconstructions
‘Gynoecium '
Only two specimens are preserved in external view. It is possible that
these are immature fruit retaining a tough protective outer skin. The holo-
type shows a clear peduncle scar ca 1 mm in diameter. It is probable that
Lindtheca gynoecia occurred in a compound strobilus similar to that of
Carnoconites.
The external architecture of the quilted, hackysacky-like, ovulate
‘gynoecium’, tf. 1 opposite, is based on the clearly preserved holotype
(PRE/F/21431a,b). A second specimen [PRE/F/21 140a,b, pi. 1 40( 1 —4)]
shows an inner view with the cell outlines preserved, but not the sutured
quilting. The internal organisation is based on information from most of the
remaining specimens. PRE/F/21432a,b, pi. 140(5, 6), shows numerous (ca
75) ovules (the darker ellipsoidal shapes), some of which are grouped
around a central cavity.
Ovuliferous ‘cell’
The ovules are interpreted to be in groups of five. This is supported by
the total number of ovules being ca 75 and the outer polygonal areas num-
bering ca 15(15 segments or ‘cells’ with five ovules each). The position of the
ovules is inferred from specimens PRE/F/12907, pi. 140(7), PRE/F/21079
and PRE/F/21 080a,b. In these cross section views the ovuliferous ‘cells’
(prism segments) are seen as triangular wedges with a darker area at the
narrow end towards the central receptacle area.
A problem with this interpretation is that the ovules are not all found
near the inferred core area. It is possible that they became spread out when
squashed prior to or during fossilisation.
Lindtheca hackysackia J.M.And. & H.M.And., sp. nov.
Holotype
Specimen'. PRE/F/21431a,b; pi. 139(1-6).
Assemblage (TC): Aas 411 Dic/Sph, Aasvoelberg.
Preserx'ation: complete ‘gynoecium’, clearly showing the outer architec-
ture and fine cellular details, part and counterpart; impression in thin-
ly laminated, strongly baked, yellowish grey shale with very good
cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 16 individuals; all detached (without peduncle), variously
complete and squashed ‘gynoecia’; the holotype and one other frag-
ment show the head in outer aspect, while the remaining 14 specimens
expose inner views of the structure with its numerous ovules (pis 139,
140).
Specific diagnosis— as for genus.
Specific characters— as for genus.
Etymology
hackysackia— 'with reference to the small, leathery, hackysacky ball (used
in a USA ball game) to which the ovulate ‘gynoecium’ of this species
bears a striking resemblance.
Comment & comparison— as for genus.
Male pentoxylalean strobili
Sahnia (Lower Cretaceous, Rajmahal Hills, India), the male pentoxy-
lalean strobilus, is branched and bears pedicellate spherical (ca 0.4
mm diam.) microsporangia; the nature of dehiscence is not known.
Pollen is apparently known only from permineralised material (Bose
et al. 1985).
Specimens of Sahnia are rare in the fossil record:
• Drinnan & Chambers ( 1 986): Koonwarra Lake deposit. Whitelaw
road cutting, Victoria, mid-Cretaceous; three mature and six
immature strobili recorded; several microsporangia illustrated.
• White (1981, 1986); Talbragar Fish Beds, latest Jurassic, New
South Wales; two isolated strobili.
• Vishnu-Mittre (1953), Bose et al. (1985); Nipania, Rajmahal Hills,
Bihar, India. Upper Jurassic to Lower Cretaceous; (?)two per-
mineralised specimens, both multiple and attached.
GONDWANA TRIASSIC, 'GEOSTRAT DISTRIBUTION
PENTOXYLALES
Lindtheca
358
d^TRELITZIA 15 (2003)
Taeniopteris Brongn. 1832
Type species
Taeniopteris vittata Brongn. 1832?
Whitby, Yorkshire, England; Middle Jurassic.
Generic concept
A form-genus for simple strap-shaped leaves with entire lamina and
lateral, parallel to occasionally forking venation at right angles to promi-
nent midrib.
Generic characters
Leaf, small to large, simple strap-shaped, tapering at base; apex acute to
obtuse; petiole short to absent, midrib prominent; venation at or close
to right angles, parallel, forking once to occasionally twice, very rarely
anastomosing.
Cuticle: see And. & And. (1989, p. 364); this vol., tf. 1 below.
Etymology
Taeniopteris— taenia (Lat.), ribbon; pteris (Lat.), fern.
Global range: numerous spp.. Pangaea, U. C. to Recent.
Gondwana Triassic occurrence
Frequency (F): 32 degree squares (of the 84 across Gondwana).
Ubiquity (U): 5 continents (of 5 comprising Gondwana).
Diversity (D): 10 foliage species.
Abundance (A): 2% (the norm as in Molteno TCs).
Longevity (L): 20 myrs (Spathian to early Norian).
Colonisation success: FUDAL rating 32/5/10/2/20 = 69.
Intermediate success (Grade 3); Taeniopteris was the 4th most promi-
nent genus in the Gondwana Triassic; it was frequent, diverse, ubiqui-
tous, widespread and long-lived, but lacking in abundance.
Endemism: of the 10 described Gondwana Triassic species, 6 are more or
less widespread, 3 are basin endemics, and 1 is a single-assemblage
endemic.
Molteno occurrence
Frequency (F): 38 TCs (of 100 sampled in the Molteno).
Diversity (D): 8 species.
Abundance (A): abundant (10%) in 3 TCs; common (3-5%) in 3 TCs;
occasional (1-2%) in 11 TCs; <1% in the other 21 TCs.
Habit: possibly woody shrubs to large trees, as suggested for T. homeri-
folius from the Molteno.
Preferred habitat: nearly ubiquitous in Dicroidium riparian forest (8 of 10
TCs), less frequent in open woodland ( 19 of 35 TCs) and lake-margin
woodland (5 of 10 TCs).
Affiliated organs
Female strobilus: Lindtheca— Grade 3 with T. homerifolius (Kin. reinf.,
Mut. occ.).
Male strobilus: unknown.
from And. & And., 1989
Classification & comparison
Suprageneric classification
While Taeniopteris is generally regarded as a form-genus, the affilia-
tion (Grade 3) of the Molteno species, T. homerifolius. with Lindtheca , sug-
gests that some forms of Taeniopteris may be placed in the Pentoxylales.
A similar case applies with regard to Doratophyllum Harris (1932).
This Taeniopteris-hke leaf has been found associated with the megasporo-
phyll Palaeocycas. As these genera show cuticular correspondence. Florin
(1933) erected the name Bjuvia for their combination.
Intergeneric comparison
Gondwana Triassic cycadopsid genera— The four cycadopsid genera
(pp. 140-146) described from the Molteno all have divided laminae to
some degree. They are quite distinct from Taeniopteris.
Other cycadopsid genera with similar leaf morphologies but not
recorded from the Molteno are: Nilssonia, Nilssoniopteris and Macro-
taeniopteris (see And. & And. 1989, p. 362). These all have characters that
distinguish them from Taeniopteris.
Pentoxylalean genera— Nipaniophyllum Sahni (1948) is another
Taeniopteris- like leaf. It has been found associated with cones and stems
and has been placed in the order Pentoxylales (Sahni 1948; Stewart &
Rothwell 1993).
Gnetopsid genera— Two Gondwana Triassic genera with taeniopteroid
venation, Yabeiella Oishi (1931), with a distinct marginal vein, and
Jungites , were placed by us (And. & And. 1989) in the Gnetopsida on the
basis of their very distinctive cuticle.
assemblages
(taphocoenosis)
Taeniopteris
T. cylomelformis
” fissiformis
” parvilocus
” anavolans
” cetilocus
” homerifolius
” yabeiformis
” sp.A (Kon 211*)
” spp. indet.
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2
J .1 -fl J _l _
-
Vin 111
Die odo
4
-i . -j .1 .1 -1 4: -1 -
-
Ela 111
Die odo
1
-1 -i 3
1 -1 -1 -1 -1 -
Kra 111
Die odo
5
^ -m 91 -; -! -
-
Lut 311
Hei elo
23
-L-! 23!
J J J J J
-
” 211
Sph pon
10
-i -i -10 -| -
-
Tin 121
Sph 2spp
2
- ! -I - -1 -) 2 -i -' -
-
Wal 111
Die odo
3
- io -;
3 -1 -1 -1 -1
-
Kon 222
Die odo
4
-1 - -!
2; -! 2; -! -i -
-
” 211
Ast 2spp
10
_ j _ 1 _ l
-lip: -
-
" 111
Die odo
1
.1 .1 . - 2 -
-
Pen 321
Dic/Ris
5
-i -i - : 5 -l -l -1 -l -
-
" 421
Die odo
2
-1 -! -i
2 -i -1 -1 -
-
Kle 111
Hei/Dic
5
r
-j-|D
Kul 111
Sph pon
1
j t
1 1 1 1 1
Kap 111
Dic/Ris
3
J -! -j
2 -! -! -! 1
-
Nuw 111
Die zub
1
-1 -|l .! -1 -1 -1 -1 -
Mor 111
Die zub
1
A
-1 -1 1: -1 "I -1 “I "I -
” ” Die odo
2
o
3; -i -, -: -
Mak 111
Die odo
10
-: - io
j j j J _i
-
Maz 211
Hei/Dic
1
-i 1
_i j j J _i
Hla 212
Die 3spp
5
-i TJ 2:
.1 3, .1 .1 -
-
” 213
Die elo
6
-1 - | -i
.! _! .
-
Umk 111
Die 2spp
55
2: 2 , -
- 50 -1 7! -! -
-
Inj 111
Die odo
4
- -1 4
J J J J J
-
” 211
Die dub
1
- - 1 - - - - - -
San 111
Die era
2
-. 2<
2 -1 2 -1 -
-
Mng 111
Die 2spp
2
2 -1 -1 -1 -1 -
-
Mat 111
Die dub
2
-1 -: 2:10; 3: -; -! -
-
Gol 111
Die dub
1
-! 1
1 1 1 1 1
Lit 111
Dic/Hei
78
-! -1 -; 20: 50 5' 3: -! -
-
Aas 111
Hei elo
2
-1 -| -1
2- -! -i -1 -1 -
-
” 411
Dic/Sph
1
A
-1 "I -1 -1 1 -1 "I "
16
Total TCs
38
* 1 1
1i 4 17'
1
18 3 13 4 1j 1
1
Tab. 62. Taeniopteris, Molteno occurrence
Taeniopteris
PENTOXYLALES
(§?'. TRELITZIA 15 (2003)
359
PENTOXYLALES
Taeniopteris
360
(3p'. TRELITZIA 15
(2003)
Lindtheca hackysackia
pi. 139
PENTOXYLALES
361
^/tRELITZIA 15 (2003)
PRE/F/2 1140a
PRE/F/2 1140a
PRE/F/21432b
PRE/F/21432b
PRE/F/21140b
Aasvoelberg
(Aas411 Dic/Sph)
PRE/F/21140a
PENTOXYLALES
pi. 140
Lindtheca hackeysackia
362
(§^r. TRELITZIA 15 (2003)
GNETOPSIDA Pilg. & Melch. 1954
NATALIGMALES J.M.And. & H.M.And., ord. nov.
NATALIGMACEAE J.M.And. & H.M.And., fam. nov.
Nataligma J.M.And. & H.M.And., gen. nov.
Type species
Nataligma dutoitii J.M.And. & H.M.And., sp. nov.
Umkomaas Valley, Karoo Basin. S. Africa, Carnian, Triassic.
Generic diagnosis
A stem-gnetopsid ovulate compound strobilus comprising a series of
whorls of small whorled ovate strobili (cones) borne on elongated slender
axes.
Generic characters
Compound strobilus: primary axis linear, gracile (ca 2 mm diam.), smooth;
strobili in widely spaced whorls (up to 50 mm apart) of ca 8 units
along axis.
Strobilus: ovoid, compact, cone-like, small (ca 5x8 mm); axis gracile,
long (ca 15-20 mm), curving upwards; megasporophylls numerous, in
ca 6 whorls (of up to 10 units) per cone.
Megasporophyll: apparently a simple scale; distal laminae rounded, imper-
fectly demarcated and forming a continuous shell; possible trichomes
visible as 2 or 3 linear parallel to spreading marks per lamina, and
more evidently extending from distal end of cone; peduncle gracile.
Ovule: unknown.
Etymology
Nataligma— an intriguing enigma from the province of Natal.
Global range: 1 sp., Gondwana, Tr. (CRN).
First & last: Molteno Fm.
Gondwana Triassic occurrence
SAf— Karoo Basin, one TC (5 indivs).
Molteno occurrence
Frequency (F): 1 TC (of 100 sampled in the Molteno).
Diversity (D): 1 species.
Abundance (A): 4 indivs; extremely rare.
Umk 1 1 1 Die 2spp: 4 indivs in 400 man-hours (1 per 8 man-days), extremely rare
Affiliated organs
Male strobilus: unknown.
Foliage: Gontriglossa— Grade 2 (Mut. occ., Mor. cor.).
Classification & comparison
Suprageneric classification (Nataligmaceae/Nataligmales)
We originally placed this compound strobilus in the Sphenophyta
(And. & And. 1983. pi. 21, fig. 1 ) based on the whorls of cones along the
axis. However, the Molteno axis lacks the striations generally characteris-
ing sphenophyte stems. The individual cones of Nataligma do somewhat
resemble Echinostachys cones which have been found attached to
Schizoneura stems with leaves coming from the same whorl. However, the
cones of Echinostachys show distinct scales in outer view. Their detailed
structure is now known (Grauvogel-Stamm 1978; Grauvogel-Stamm &
Ash 1999).
Amongst the gymnosperms, the only groups of which we are aware
that bear whorled fruiting structures are the Gnetopsida (extant) and the
Fredlindiales (Gondwana Triassic). Is there a shared plesiomorphic character
involved, or is the similarity an expression of convergent evolution? The
small individual cones of Nataligma are certainly unlike anything within
these two groups.
Nataligma is included here, alongside the Fraxinopsiales, primarily on
the basis of the cuticular similarities between their foliage affiliates (Grade
4 reliability in the latter, but only Grade 2 in the former). The pair are
placed in the class Gnetopsida in view of their shared, possibly symple-
siomorphic, whorled character. Fraxinopsis in turn has a possible link to
Dechellyia, Late Triassic, USA (see box, p. 373), which has been classified
in the Gnetopsida (Ash 1972; Crane 1988). Nataligma is placed in the
new order Nataligmales and new family Nataligmaceae within the class
Gnetopsida.
Intergeneric comparison (Gondwana Triassic): nothing similar.
A. dutoitii
based on
PRE/F/6857 & PRE/F/6858
pi. 141(1,2,6,7)
Umk 111
Reconstructions
The holotype, pi. 14 1 ( 1—4), shows two whorls of strobili. The primary
axis is of similar width for 88 mm, an indication that the total length was
probably far greater, A second specimen, with one whorl, pi. 141(5-8),
could be a part of the holotype as it matches in size— but the cleavage does
not fit owing to missing rock matrix. We have drawn the compound stro-
bilus with three whorls, leaving the proximal and distal ends incomplete.
The nature of the individual strobilus or cone is evident in
PRE/F/6858a,b, pi. 142(1-3), where six whorls are visible. These whorls of
pedicellate scales (megasporophylls) blend into the glossy carbonaceous
matrix. Two or three fine linear striations which may represent trichomes
are seen repeated on each of the ill-defined distal laminae [PKE/F/6857a,b,
pi. 142(4-6)]. On another isolated strobilus, PRE/F/6859b, a pair of possi-
ble trichomes can be seen to arise from the apex of the cone, pi. 142(7, 8),
as does a single hair from the left side of the strobilus, pi. 142(9).
Nataligma
NATALIGMALES
TRELITZIA 15 (2003)
363
Evidence for affiliation of organs
Mutual occurrence
Nataligma is found only at Umk 111, which also yields Gontriglossa
verticillata (Tabs 25, 26).
Morphological correspondence
The primary axis (stem) of this compound strobilus is very similar to
the stem of Gontriglossa verticillata (And. & And. 1989), which bears
leaves in apparent whorls (composed of opposite fascicles) at regularly
spaced nodes ( ca 100 mm apart). A good collection of G. verticillata ,
including three stems with leaves attached, is known from the same TC
(Umk 111) as Nataligma. On the basis of similar morphology— a stem with
leaves or cones in whorls— we place Nataligma as an affiliate (Grade 2) of
Gontriglossa verticillata.
Cuticles
Potential sample : Umk 111,4 indivs.
Macerated (this work): 2 indivs (8 samples).
Preservation grade'. Grade 1 .
Diagnostic characters: —
Comment: Microscopic details remain unknown for this Molteno cone, the
eight carbonaceous samples macerated having yielded no definable
morphology. The cuticle is apparently quite altered.
Significance: of no aid in classification or affiliation.
Nataligma dutoitii J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/6857a,b; pis 141(1-4), 142(4—6).
Assemblage (TC): Umk 1 1 1 Die 2spp, Umkomaas Valley.
Preservation: middle section of strobilus, part and counterpart; compression
in thinly laminated, carbonaceous (good cuticle), moderately baked,
dark grey shale with good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 4 indivs (2 intact strobili, 2 isolated cones), pis 141, 142.
Specific diagnosis— as for genus.
Specific characters— as for genus.
Eponymy
dutoitii— in honour of Alexander du Toit, the greatest of South African
geologists and a pioneer in the study of the Molteno palaeoflora.
Comment & comparison— as for genus.
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
NATALIGMALES
Nataligma
364
d^TRELITZIA 15 (2003)
Goiltriglossa J.M.And. & H.M.And. 1989
Type species
Gontriglossa verticillata (Thomas 1958) J.M.And. & H.M.And. 1989.
Umkomaas Valley, Karoo Basin, S. Africa; Camian, Triassic.
Generic diagnosis
A stem-gnetopsid vegetative genus comprising long shoots bearing
opposite fascicles of narrowly elliptic leaves, with frequently anastomosing
open-mesh arching venation, at regularly spaced nodes.
Generic characters
Foliage shoot: leaves apparently attached in whorls comprising opposite
fascicles of 3 leaves at regularly spaced nodes (ca 100 mm apart);
shoot slender (4-6 mm diam.), with indistinct longitudinal striations.
Leaf, medium (110 X 25 mm); petiole stout, moderately long; lamina nar-
rowly elliptic, margin entire to irregularly wavy, apex obtuse; midrib
distinct, veins at steep angle (ca 50°) from horizontal, arching strong-
ly to margin, frequently anastomosing, mesh open, length to width
ratio 10; 1 near midrib and decreasing strongly to margin.
Cuticle: see And. & And. (1989, p. 505); this vol., tf. 4 opposite.
Etymology
Gontriglossa— a combination word referring to Glossopteris- like leaves
from the Gondwana Triassic.
Global range: 1 sp., Gondwana, U. Tr. (LAD-CRN).
First: Glossopteris (Gontriglossa) grandis (Holmes 1992); Basin Creek Fm.,
Nymboida, N.S.W., Australia.
Last: the Molteno species.
Gondwana Triassic occurrence
Frequency (F): 10 degree squares (of the 84 across Gondwana).
Ubiquity (U): 4 continents (of 5 comprising Gondwana).
Diversity (D): 1 foliage species.
Abundance (A): 1% (the norm as in Molteno TCs).
Longevity (L): 9 myrs (Ladian to Camian).
Colonisation success: FUDAL rating 10/4/1/1/9 = 25.
Limited success (Grade 2); Gontriglossa was the 13th most prominent
genus in the Gondwana Triassic; it was moderately frequent, ubiqui-
tous, long-lived and common, but markedly lacking in diversity.
Endemism: The single species, G. verticillata, had a wide distribution, being
known from four Gondwana continents.
Molteno occurrence
Frequency (F): 8 TCs (of 100 sampled in the Molteno).
Diversity (D): 1 species.
Abundance (A): common (5%) in 1 TC; occasional ( 1-2%) in 2 TCs; rare
to extremely rare (<1%) in 5 TCs.
Habit: possibly a slender herbaceous pioneer.
Preferred habitat: frequent in the Dicroidium riparian forest (6 of 10 TCs);
also found in fem/horsetail meadows (wetlands).
Affiliated organs
Female strobilus: Nataligma— Grade 2 (Mut. occ., Mor. cor.).
Male strobilus: unknown.
Classification & comparison
Intergeneric comparison
Three historical genera in particular, all erected in the 19th century,
have been repeatedly employed in the Gondwana Triassic literature for
Gontriglossa verticillata leaves: Glossopteris, Sagenopteris and Anthrophy-
opsis (And. & And. 1989, pp. 504, 505).
Glossopteris Brongn. 1828 (dominant in the Gondwana Permian) — is
superficially indistinguishable from Gontriglossa, but the cuticle and mode
of attachment are different.
Sagenopteris Presl 1838 (the foliage of the Caytonia plant) — was
widespread in Laurasia and ranged from the Upper Triassic to the Upper
Cretaceous (Crane 1985). G. verticillata may be confused with the indi-
vidual leaflets of Sagenopteris, but the latter is a compound, superficially
palmate leaf, with two pairs of readily dehiscing, sessile leaflets, attached
apically on a long petiole, and it has a cuticle with anomocytic stomata and
cells of a regular shape with straight to sinuous walls (Harris 1964; Crane
1985).
Anthrophyopsis Nathorst 1878 (Rhaetic of Sweden and Greenland) —
has cuticle which is also quite different from G. verticillata. Its mode of
attachment and reproductive organs are unknown.
Among Gondwana Triassic foliage genera, G. verticillata has much in
common with Yabeiella (widespread and common) and Jungites (exclusive
to the Molteno). Each shows anastomosing venation to lesser or greater
extent and their cuticles are similar in having amorphously shaped non-
papillate cells with strongly meandering walls. While the cuticles of
Yabeiella and Jungites are very close, that of G. verticillata differs in hav-
ing paracytic stomata and complex digitately amorphous cell outlines. It
seems likely that the three genera, together with Graciliglossa and
Cetiglossa, comprise a natural group or clade. here included in the
Gnetopsida.
Holmes (1992) placed certain Australian Triassic leaves in the genus
Glossopteris— while emphasising that Glossopteris was used as a form
genus and that no relationship with Permian Glossopteris was inferred.
Glossopteris grandis (Walkom) Holmes has whorls of up to 10 leaves
which appear, individually, to be closely similar to Gontriglossa verticilla-
ta. However, the form of attachment to a stem is not clear, so close rela-
tionship cannot be made with the Molteno material.
Gontriglossa
NATALIGMALES
d?TRELITZIA 15 (2003)
365
GONDWANA TRIASSIC, 'GEOSTRAT DISTRIBUTION
NATALIGMALES
Gontriglossa
366
d/TRELITZIA 15 (2003)
IsLi j£SH8&38 jt.£s
PRE/F/6857b-'
Holotype
PRE/F/6857b ,
_ . VT1L
PRE/F/6858b
PRE/F/6857b >;
\ PRE/F/6857b
>■■■11 iiiliiii w It > i
Umkomaas Valley
(Umk 1 1 1 Die 2spp)
Jp
PRE/F/6858b
I PRE/F/6858a
X5 PRE/F/6858a
Nataligma dutoitii
pi. 141
NATALIGMALES
367
^/trELITZIA 15 (2003)
NATALIGMALES pi. 142 Nataligma dutoitii
368
^/trelitzia 15 (2003)
GNETOPSIDA piig. & Melch. 1954
INCERTAE SEDIS order
INCERTAE SEDIS family
Graciliglossa J.M.And. & H.M.And., gen. nov.
Type species
Gontriglossa hilaryjanea J.M.And. & H.M.And. 1989.
Umkomaas Valley, Karoo Basin, S. Africa; Camian, Triassic.
Generic diagnosis
A stem-gnetopsid leaf of small gracile form, oblong-lanceolate shape
with cordate base and entire margin, and frequently anastomosing venation
with open mesh at steep angle to margin.
Generic characters
Leaf, small (60 X 15 mm), gracile, oblong-lanceolate, base cordate-sagittate,
apex obtuse, lamina margin entire; petiole gracile, seemingly short;
midrib distinct; veins at very steep angle (ca 70-80°) from horizontal,
not arching to margin, frequently anastomosing, mesh open, length to
width ration 6: 1 near midrib and decreasing strongly to margin, with
distinctive marginal vein parallel to cordate base.
Cuticle : unknown.
Graciliglossa hilaryjanea J.M.And. & H.M.And., comb. nov.
Holotype
Specimen : BP/2/590; And. & And. [1989, pi. 309(3, 14)].
Assemblage (TC): Umk 111 Die 2spp, Umkomaas Valley.
Preservation', nearly complete leaf, without counterpart; compression in
thinly laminated, carbonaceous (good cuticle), moderately baked, dark
grey shale with good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 13 indivs [And. & And. 1989, pis 309(1-22), 310(1-11)].
Sister palaeodemes— none.
Specific diagnosis— as for genus.
Specific characters— as for genus.
Eponymy
hilaryjanea— for our daughter Hilary Jane, who is a gracile child and who
has accompanied us on several collecting trips.
Comment & comparison
No similar leaf is known from the Gondwana Triassic.
Etymology
Graciliglossa— gracilis (Lat.), gracile, thin, slender; glossa (Gr.), tongue;
referring to the gracile, tongue-shaped leaf.
Global range: 1 sp., Gondwana, U. Tr. (CRN).
First & last: the Molteno species described here.
Gondwana Triassic occurrence
Frequency (F): 1 degree square (of the 84 across Gondwana).
Ubiquity (U): 1 continent (of 5 comprising Gondwana).
Diversity (D): 1 foliage species.
Abundance (A): <1% (as recorded in the Molteno).
Longevity (L): 1 myr (Lower Camian).
Colonisation success: FUDAL rating 1/1/1/— /1= 4.
Minimum success (Grade 1); Graciliglossa was an extremely minor
component of the Molteno flora.
Endemism: the single species, G. hilaryjanea , as known, is a single-assemblage
endemic.
Molteno occurrence
Frequency (F): 1 TC (of 100 sampled in the Molteno).
Diversity (D): 1 species.
Abundance (A): very rare (<1%) in the 1 TC.
Habit: possibly a slender, herbaceous pioneer.
Preferred habitat: Dicroidium riparian forest.
Affiliated organs: unknown.
Classification & comparison
We have transferred this leaf to a new genus distinct from Gontriglossa
because of its small size, gracile axis and petiole, cordate base with marginal
vein and the lack of specimens showing the attachment of leaves as found
in that taxon (see further under Cetiglossa, p. 369).
No similar leaf is known from the Gondwana Triassic, but in the
Permian certain Glossopteris leaves have a cordate base and open anasto-
mosing veins.
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
Graciliglossa
GNETOPSIDA/INCERTAE
^/tRELITZIA 15 (2003)
369
GNETOPSIDA Pilg. & Melch. 1954
INCERTAE SEDIS order
INCERTAE SEDIS family
Cetiglossa J.M.And. & H.M.And., gen nov.
Type species
Gontriglossa balaena J.M.And. & H.M.And. 1989.
Umkomaas Valley, Karoo Basin, S. Africa; Camian, Triassic.
Generic diagnosis
A stem-gnetopsid leaf of large size, broadly elliptic shape with irregu-
larly crenulate margin, and frequently anastomosing venation with elongate
parallel mesh at moderate angle to margin.
Generic characters
Leaf, large (up to >110 mm long), broadly elliptic, apex probably obtuse;
lamina margin irregularly crenulate; petiole unknown; midrib distinct,
veins at moderate angle (ca 40-45°) from horizontal, very faintly sin-
uous to margin, frequently anastomosing, mesh elongate, length to
width ratio ca 15:1 near midrib and decreasing strongly to margin.
Cuticle : unknown.
Etymology
Cetiglossa— cetis (Lat. ), whale; glossa (Gr.), tongue; with reference to the
tongue-shaped leaf from the Umkomaas locality.
Cetiglossa balaena (J.M.And. & H.M.And. 1989) J.M.And.
& H.M.And., comb. nov.
Holotype
Specimen : PRE/F/750; And. & And. [1989, pi. 308(4, 6-9)].
Assemblage (TC): Umk 111 Die 2spp, Umkomaas Valley.
Preservation', incomplete leaf, without counterpart; compression in thinly
laminated, carbonaceous (good cuticle), moderately baked, dark grey
shale with good cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 3 indivs (And. & And. 1989, pi. 308).
Species diagnosis— as for genus.
Species description — as for genus.
Etymology
balaenus (Lat.)— whale, after the type locality Umkomaas (meaning cow
or whale in the local Zulu language).
Comment & comparison
Glossopteris nymboidensis Holmes (1992), from the Middle Triassic
Basin Creek Fm., Nymboida Coal Measures, Australia, is close to C. bal-
aena, but differs in the still narrower, more elongate venation mesh and in
the decurrent lateral veins which leave the midrib at an acute angle, then
arch and continue straight and parallel towards the margin at ca 15°.
Global range: 1 sp.. Gondwana, U. Tr. (CRN).
First & last: the Molteno species described here.
Gondwana Triassic occurrence
Frequency (F): 1 degree square (of the 84 across Gondwana)
Ubiquity (U): 1 continent (of 5 comprising Gondwana).
Diversity (D): 1 foliage species.
Abundance (A): <1% (as recorded in the Molteno).
Longevity (L): 1 myr (Lower Camian).
Colonisation success: FUDAL rating 1/1/1/ — /I =4.
Minimum success (Grade 1); Cetiglossa was an extremely
minor component of the Molteno flora.
Endemism: the single species, C. balaena, as known, is a
single-assemblage endemic.
Molteno occurrence
Frequency (F): 1 TC (of 100 sampled in the Molteno).
Diversity (D): 1 species.
Abundance (A): extremely rare (<1%) in the single TC.
Habit: possibly a slender herbaceous pioneer.
Preferred habitat: Dicroidium riparian forest.
Affiliated organs: unknown.
Comparison & classification
Intergeneric comparison
In And & And. (1989), the three species, Cetiglossa balaena, Gracili-
glossa hilaryjanea and Gontriglossa verticillata, were all included in the
genus Gontriglossa. These are now placed in three distinct genera, reflect-
ing a more natural classification, we believe. The changes are based on the
following reasoning:
1 ) Although all three species have clear midribs and reticulate venation,
they differ particularly widely in size, leaf shape and nature of the
anastomosing veins.
2) Gontriglossa verticillata has yielded very characteristic Grade 3 cuti-
cle, while Graciliglossa hilaryjanea and C. balaena have yielded only
Grade 1 cuticle without interpretable structures. Were the three species
of the same genus, one might expect all to yield cuticle of similar quality.
3) While Gontriglossa verticillata has been found frequently with leaves
attached to a stem, the other taxa are unknown in this state. This sug-
gests modes of attachment that might have been significantly different.
Subject to confirmation, based on a good deal more information than
the foliage alone, we place Cetiglossa and Graciliglossa in separate fami-
lies but a single order (all unnamed), probably distinct from the Natalig-
males (see pp. 21. 55).
Santaecruzia (Gnaedinger & Herbst 1998) was based on Gondwana
material from the Upper Triassic of Argentina. The genus has much finer,
closer-spaced anastomosing venation than Cetiglossa and the veins emerge
at an angle of 90° to the midrib.
INCERTAE/GNETOPSIDA
Cetiglossa
370
C^T RELITZIA 15 (2003)
GNETOPSIDA Pilg. & Melch. 1954
FRAXINOPSIALES J.M.And. & H.M.And., ord. nov.
FRAXINOPSIACEAE J.M.And. & H.M.And.. fam. nov.
Fraxinopsis Wieland 1929
Type species
Fraxinopsis minor Wieland 1929.
Minas de Petroleo, Cacheuta Basin, South America. Potrerillos Fm., L.
Camian, Triassic.
Generic concept
A stem-gnetopsid dispersed seed (strobilus unknown), comprising a
small seed body with longitudinal groove, and a large, elongate leaf-like
wing with parallel veins that may bifurcate and anastomose.
Generic characters
Strobilus : unknown.
Megasporophyll : unknown.
Seed: winged, small to large (ca 1 2— >50 mm long); seed body relatively
small, elliptical, with a distinct longitudinal groove, glabrous to dense-
ly hairy, with incipient to pronounced distal auricular lobes; wing leaf-
like, relatively large, elongate to oval-elliptic, 2-5 times the length of
the seed body, glabrous to hirsute; veins clearly developed, ca 6-16 at
widest point, parallel and rarely to commonly bifurcating and anasto-
mosing.
Etymology
Fraxinopsis— after the extant genus Fraxinus, whose seeds Wieland (1929)
considered similar to his fossil material.
Global range: several spp., Gondwana, Tr. (CRN).
First: Fraxinopsis major (Jones & De Jersey 1947); Tivoli Stage, Ipswich
CM, Queensland. Australia.
Last: Fraxinopsis andium: Molteno Fm.
Gondwana Triassic occurrence
SAm— N. Argentina; Cacheuta Basin, 4 Iocs (many indivs reported);
Barreal-Hilario Basin; 2 Iocs (6 indivs illustr.).
SAf— Karoo Basin (Molteno Fm.); 17 TCs (292 curated indivs).
Aus— Queensland; Ipswich Coalfield, 11 Iocs (many indivs reported).
Molteno occurrence
Frequency (F): 18 TCs (of 100 sampled in the Molteno).
Diversity (D): 3 species.
Abundance (A): 306 indivs total; rare to extremely rare in top 9 TCs.
Kra 311 Die odo: >35 indivs in 13 man-hrs cleaving (>10 per 1 man-day) rare
Maz 211 Hei/Dic:
Kap 111 Dic/Ris:
San 111 Die era:
63 ”
26 ”
10 "
” 85
” 65
” 30
” (
” (
” (
8 ” 1 ” ) ”
5 ” 1 ” ) ”
3 ” 1 ” ) ”
Wal 111 Die odo:
12 ”
” 50
(>
1 ” 1 ” ) v. rare
Bir 111 Sph 2spp:
47 ”
” 550
” (>
1 ” 1 ” ) ”
Aas 411 Dic/Sph:
44 "
” 512
” (>
1 .. 1 -• ) »
Umk 1 1 1 Die 2spp:
14 ”
'• 400
” (
1 ” 3 ” ) ”
Lit 111 Dic/Hei:
10 ”
” 550
” (
1 ” 5 ” ) ex. rare
The above nine TCs (of the 1 8 yielding Fraxinopsis) are covered in the
photographic plates. For the TCs (Maz 211, Bir 111 and Aas 411) with 30
or more curated individuals, the recorded abundance is conservative as
only the better preserved specimens have been retained (see Tab. 64).
Affiliated organs
Male strobilus: unknown.
Foliage: Yabeiella— Grade 4 (Cut. cor., Mut. occ.).
Classification & comparison
Suprageneric classification (Fraxinopsiaceae/Fraxinopsiales)
Fraxinopsis shows similarities, as noted by Wieland (1929), to both
Fraxinus (extant angiosperm) and Cycadocarpidium (Triassic pinopsid),
but apparently bears no phylogenetic relationship to either. It affiliates
repeatedly with the leaf genus Yabeiella in South Africa, South America
and Australia. Yabeiella is a distinctive leaf, with basic taeniopteroid vena-
tion. irregular anastomoses and clear marginal vein. It is restricted to the
Gondwana Triassic and is entirely unlike that of Fraxinus or of
Podozamites which is affiliated with Cycadocarpidium. The venation and
cuticular features (And. & And. 1989, p. 472) of Yabeiella suggest links
with the Gnetopsida. The Fraxinopsis/Yabeiella plant, all considered, is
placed in the new order Fraxinopsiales and new family Fraxinopsiaceae.
Reconstructions
Only minor reconstruction clarifying venation, auricular lobes or hir-
suteness has been attempted in the numerous sketches of the seeds. No
information is available on how the seed was attached to the plant.
Fraxinopsis
FRAXINOPSIALES
TRELITZIA 15 (2003)
371
Gondwana Triassic occurrence (elaborated)
Fraxinopsis is well known and common from three disjunct regions of
Gondwana— N. Argentina, the Karoo Basin of South Africa and the
Clarence-Moreton Basin, Queensland. It is clearly more frequent and abun-
dant in these regions than is reflected in the hypodigm alone. Jones & De
Jersey (1947), for instance, report Fraxinopsis (without giving any abun-
dance data) from 10 Ipswich Coalfield localities— Kholo, Tivoli and
Cooneana Stages of the Ipswich Series, while Jain & Delevoryas (1967)
record 19 specimens in the Wieland collection from the Minas de Petroleo
locality, Cacheuta Basin.
South America (N. Argentina)
1929, Wieland: Fraxinopsis major and F. minor ; f5 (a, b); new genus and two species
based on ca 10-20 specimens, Minas de Petroleo, Cacheuta Basin, N. Argentina;
Potrerillos Fm. (Camian, U. Triassic).
1931, Oishi; Fraxinopsis minor, pi. 26(1, la); a single specimen from near Minas de
Petroleo; Potrerillos Fm.
1941a, Frenguelli: Cycadocarpidium andium; pi. 1-3, tf. 1-3; a new species based on
five illustrated specimens. Estratos con Estheria, Cacheuta Basin; Cacheuta Fm.
(Camian, U. Triassic).
1941b, Frenguelli: repeats after Wieland 1929.
1944b, Frenguelli: Cycadocarpidium major, C. minor and C. andium : pi. 1-4 (ca 20
specimens); locality behind old YPF building, Cacheuta Basin; Potrerillos Fm.
1963, Bonetti: Cycadocarpidium andium, pi. 39(1—4), pi. 40(1, 2); 6 specimens illus-
trated; Punto 20, 21, Barreal-Hilario Basin, N. Argentina; Cortaderita Fm. (Camian,
U. Triassic).
1967, Jain & Delevoryas: Fraxinopsis major and F. minor, pi. 97(9-14), six specimens
illustrated, 19 specimens recorded; Minas de Petroleo (site and collection as for
Wieland 1929).
1969, Menendez: Cycadocarpidium andium, pi. on p. 535(1 ); repeat after Frenguelli
1941a.
1995, Ganuza et at.: Cycadocarpidium andium and C. majus. pi. 2(g, h), two specimens
illustrated and recorded; Paso Flores region and Fm.; here identified as belong-
ing to F. cornicordis.
Australia (Queensland, Clarence-Moreton Basin)
1947, Jones & De Jersey: Fraxinopsis major, p. 54, tfs 49, 50; specimens without
abundance data from 10 localities, Ipswich Coalfield, Clarence-Moreton Basin,
Queensland; Kholo, Tivoli & Cooneana Stages, Ipswich Series (Camian, U.
Triassic).
1965, Hill et al.\ Fraxinopsis major, pi. T6(6); a single specimen illustrated from
Petrie's Quarry, Albion, Ipswich Coalfield; Tingalpa Fm., Ipswich Series.
Tab. 63
FRAXINOPSIS HYPODIGM, Gondwana Triassic occurrence
Species
Molteno
Other
2
2 "6
c 2 o
§ 3 o
1 -a f
F. minor
F. major
F. spp. Indet
AUTHOR
SUBREGION
FORMATION
LOCALITY
NAME
Indivs ILLUSTRATION
C 3
ro ro
u: u:
O
o
u:
SOUTH AMERICA
i
i
1929 Wieland
Cacheuta NA4
24 Potrerillos
Minas de Petroleo
Fraxinopsis major
1 f 5(a)
- 1 -
-
- 1 -
|
” minor
2 f 5(b)
- ; -
-
2 - -
1931 Oishi
” | "
” |
1 pi 26(1, la)
-
1 ] -j -
1941a Frenguelli
” | "
25 Cacheuta
(Estratos con Estheria)
Cycadocarpidium andium
5 pi 1-3, tf 1-3
5 -
- 1 - 1 -
1941b ” (specimen repeated from Oishi 1931, Frenguelli 1941b)
* 1
1 1
1944b ”
Cacheuta NA4
24 Potrerillos
Old YPF admin. Building
Cycadocarpidium minus
8 pi 1(1-7), 4(1,2), tf 1 A
i ; -
-
8 - -
” ] ”
„ , „
” |
andium
8 pi 2(1-7), 4(2), 5(1)
8 -
- 1 -J -
.. j
” |
majus
5 pi 3(1-5), tf IB
- 1 -
-
- 5; -
1963 Bonetti
Barreal NA2
23 Cortaderita
Punto 20,21
andium
6 pi 39(1-4), 40(1,2)
6 -
- I -I -
1967 Jain & Delev.
Cacheuta NA4
24 Potrerillos
Minas de Petroleo
Fraxinopsis major
3 pi 97(9-11)
- -
-
- : 3i -
” | ..
” |
” minor
3 pi 97(12-14)
- -
-
3 - -
1969 Menendez (specimen repeated from Frenguelli 1941b)
*
j
1 995 Ganuza et at.
Paso Flores SA1
24 Paso Flores El Cahadon de Pancho
" andium
1 Pi 2(g)
- 1 -
i
” | ”
” ; ”
” |
" majus
1 Pi 2(h)
- 1 "
i
- i -i -
AUSTRALIA
[
1 1
1947 Jones & de J.
Ipswich/Esk CM5
23 Tivoli Stage
Chuwar (Iocs. 5,6)
Fraxinopsis major
2 tf 49,50
- ! -
-
-' 2i -
1965 metal.
Brisbane CM6
" Tingalpa Fm.
Petrie's Ouarry (Albion)
1 pi T6(6)
-
- ! i! -
SOUTH AFRICA
I
1
I i
1978-1999 And. & And. Molteno literature not included in this table
i I
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
FRAXINOPSIALES
Fraxinopsis
372
d^TRELITZIA 15 (2003)
Evidence for affiliation of organs
The link between the foliage genus and the winged seed Fraxinopsis is
virtually certain (Grade 4 reliability) short of organic attachment. This is
based on repeated mutual occurrence on three Gondwana continents —
South Africa, South America and Australia— and on the close similarity of
their characteristic cuticle. Meanwhile, the male affiliate of Yabeiella
remains unknown.
Mutual occurrence
South Africa— Fraxinopsis is known from 18 of the 100 Molteno TCs
studied (Tab. 64). In the majority of cases (14 of 18) it occurs with
Yabeiella , which is known from a total of 29 TCs. At least three of the four
TCs (Bir 211, Bir 311 and Mol 111) with Fraxinopsis, but no Yabeiella,
would very likely yield the latter on more comprehensive sampling. These
three TCs all represent floodplain lake deposits dominated by the foliage
genus Sphenobaiera, as do the very well-sampled sites, Bir 111 and Aas
411, that have yielded Yabeiella fairly commonly along with Fraxinopsis
(Tab. 64).
At the remaining TCs where Yabeiella is very rare, it is not surprising
that Fraxinopsis has not been found. However, there are four TCs where
Yabeiella is fairly common ( 1 % of the total flora at Nav 111, Kon 111, Kon
222 and Hla 213) but no Fraxinopsis has been found. Three of these TCs
occur in Dicroidium open woodland habitats (Tab. 64).
South America— The genera Fraxinopsis and Yabeiella are clear affili-
ates at five of the six North Argentinean localities (Tab. 63) yielding the
winged seed.
Australia— Jones & De Jersey (1947) make special note of the ‘constant
association’ of Fraxinopsis and Yabeiella : ‘In the Ipswich Series Fraxinopsis
has only been found at localities at which species of Yabeiella occur’.
Cuticular correspondence
The cuticle of Yabeiella (p. 377, tfs 11, 12) and Fraxinopsis, with the
deeply sinuous cell walls and narrowly elliptic guard cells, is essentially
identical and quite distinct from other Molteno plants.
Cuticles
Potential sample : Lit 111, 10 indivs; Untk 111, 12 indivs.
Macerated (this work): Lit 1 1 1 , no results from three macerations (but two
peels from wing area productive).
Preservation grade: Grade 3 (fair), some features present, large pieces.
Diagnostic characters: cells amorphous, walls meandering, nonpapillate;
stomata interveinal (whether adaxial or abaxial unknown), orientation
longitudo-random; subsidiary cells anomocytic, noncutinised; guard
cells narrowly elliptic.
Comment: A distinctive feature is the elongate hairs visible along the wing
margin.
Significance:
Classification— See notes on pp. 370, 376.
Affiliations— The cuticle unequivocally supports the affiliation already
suggested by the repeated mutual occurrence of Fraxinopsis and Yabeiella.
Species nomenclature in Fraxinopsis
The particular problem encountered here in applying South American
Fraxinopsis names to Molteno material revolves around the difficult tax-
onomy/nomenclature relationship (see Hypodigm, Tab. 63).
Weiland (1929) described Fraxinopsis minor and F. major based on an
undisclosed number of specimens from the single Minas de Petroleo local-
ity, Potrerillos Fm.. Cacheuta Basin. The F. minor holotype measures 24
mm in length and shows largely parallel venation, while the F. major holo-
type is 30 mm long with more spreading dichotomous venation. The for-
mer species was apparently the more abundant.
From our understanding of over 250 Fraxinopsis specimens from 18
Molteno TCs, only one palaeodeme occurs at any one site and each shows
a marked range of morphological variation. This pattern may apply also to
the South American material. Jain & Delevoryas (1967) illustrated a further
three specimens of each species from the original Minas de Petroleo locality,
but provided no information on their spread of variation. This is a critical
point. If the collection does indeed represent a single palaeodeme with a
continuous range of variation in size, then F. major and F minor represent
a single species.
F. andium, described by Frenguelli ( 1941a), includes the holotype with
a length of 18 mm and the four additional illustrated specimens of similar
size (18-20 mm). The common and variable Molteno species (from 16
TCs) has a size range of ca 9 mm to a maximum of 24 mm. The Bir 1 1 1
reference palaeodeme ranges from ca 12-20 mm. The two additional
species from the Molteno, F. auriculata and F. cornicordis, are clearly dis-
tinct and range upwards from 32 mm in length.
We place our common Molteno species in F. andium and recognise F.
minor and F. major as distinct species until further studies are undertaken
on the South American material. The two specimens described by Ganuza
et al. (1995) as F. andium and F. majus, with lengths of 39 and 45 mm
respectively, are here both identified as F. cornicordis.
Adaptive radiation (Molteno diversity)
Palaeodemes of the foliage genus Yabeiella often show a good deal of
variation and it is difficult to demarcate species. A similar situation is
encountered in the winged seed genus Fraxinopsis, where considerable
intra- and interpalaeodeme variation occurs. Of the 18 Molteno palaeo-
demes (each from a different TC), only two are clearly distinct from the rest
and are recognised as separate species. The majority (16 TCs) fall into F.
andium originally described from Argentina.
The three species are distinguished on the basis of size, wing venation,
hirsuteness and shape of seed head. They are based on the following
TCs/reference palaeodemes, all of which occur in floodplain deposits:
F. andium— Bis 111 Sph 2spp, (Birds River); 47 indivs
Sphenobaiera closed woodland (floodplain lake); Cycle 5 (Tsomo Member)
F. auriculata — Wal 111 Die odo, (Waldeck); 12 indivs
Sphenobaiera closed woodland (floodplain lake); Cycle 2 or 3 (Indwe & Mayaputi
Member)
F. cornicordis— Kra 311 Die odo (Kraai River); 35 indivs
Dicroidium open woodland (floodplain); Cycle 3 (Mayaputi Member)
F. andium 1 F. andium 2
Fraxinopsis
FRAXINOPSIALES
c/trelitzia 15 (2003)
373
assemblages
(taphocoenosis)
Genera
Species
Molteno Cycles
Yabeiella
OFraxinopsis
F. andium
F. auriculata
F. cornicordis
Nav 111
Die odo
1 -
_ 1 1
Bir 211
Sph 2spp
1
1 ; - ! -
” 311
Hei/Sph
-i 5
5 ' -i -
" 111
Sph 2spp
40 47
47 -! -
Gre 111
Equ sp.
1 1
1 ; -
Boe 111
Lep sto
5| -
1 1
4/5
Dic/Hei
1: 1
1 -j •
Mol 111
Sph pon
-1 2
2 - ' -
Ela 111
Die odo
6
- ' - ! -
Kra 311
”
18 35
-! -i35
Lut 311
Hei elo
9 6
6 : -| -
3
” 211
Sph pon
4 -
Tin 121
Sph 2spp
1 1
1 : -
Wal 111
Die odo
20 12
-i12 -
2/3
Kon 223
1 -
” 222
1 -
_ 1 _ 1
” 211
Ast 2ssp
1 1 -
t 1
1 1
2f
" 111
Die odo
1 -
Pen 321
Dic/Ris
2 -
- 1 - ! -
Kle 111
Hei/Dic
-! 1
1 : - ! -
Kap 111
Dic/Ris
5; 26
26 - ; -
2e
Qua 111
Die odo
5 -
Maz 111
Die era
22 27
27 -
2c
’’ 211
Dic/Hei
11 63
63 > -! -
Hla 211
Die 3spp
1 . -
- ! - ; -
” 213
Die elo
1 -
Umk 111
Die 2spp
45 ' 14
14 -; -
2b
Inj 111
Die odo
2; -
- 1 - : -
San 111
Die era
2 10
10 -1 -
Mat 111
Dic/Hei
2; -
Lit 111
Die odo
6: 10
10 -i -
2a
Aas 411
Dic/Sph
60 44
44 -
1
” 511
Die elo
1! -
Total TCs
29:18
i6 1 ; 1
Total indivs
% 1 306
259 12 35
Tab. 64. FraxinopsislYabeiella,
Molteno occurrence
Beyond Gondwana Triassic
Laurasian Triassic. Dechellyia (tfs 1, 2 adjacent), a winged seed
attached to the foliage of the same name, from the Late Triassic of
southwestern United States (Ash 1972), is somewhat similar but dif-
fers in lacking a groove on the seed body and in the two veins or
thick ribs that nearly converge in the wing base. Cycadocarpidimn,
which is widespread in Laurasia, is superficially similar to
Fraxinopsis but is known to be a conifer scale belonging to the
Voltziales (And. & And. 1989, p. 423; Axsmith et al. 1997, p. 304).
Fraxinopsis has recently been recorded from the Late Triassic of
eastern United States (Axsmith et al. 1997). Their F. aquilonaris (tf.
5 adjacent) is similar to Gondwana Fraxinopsis species but differs in
lacking a groove along the seed body and in the absence of any fork-
ing or anastomoses in the wing venation. Yabeiella does not occur in
association with F. aquilonaris and no other affiliated leaf has been
suggested. It is most probable that the USA species should be placed
in a separate genus.
Other ages: Wieland (1929) originally compared Fraxinopsis with
the winged seeds of the extant angiosperm Fraxinus and suggested
that the seed had two cotyledons. However, there is no support for
this view (Oishi 1931).
FRAXINOPSIALES
Fraxinopsis
374
^TRELITZIA 15 (2003)
Fraxiliopsis andium (Freng. 1941a) J.M.And. & H.M.And.,
comb. nov.
Holotype
Specimen: 4755 Mus. de la Plata, Argentina; Frenguelli [1941b, pi. 1(1, 2)].
Assemblage: Estratos con Estheria , Cacheuta Basin, N. Argentina; Cacheuta
Fm., Camian.
Preservation: virtually complete seed; part and counterpart.
Reference palaeodeme
Assemblage (TC); Bir 111 Sph 2spp, Molteno Fm.
Specimens: 47 indivs; pi. 144(1-10).
Sister palaeodemes— 15 (best 6 listed)
Kap 111 Dic/Ris: 26 indivs; fine preservation.
San 1 1 1 Die era: 10 indivs; fine preservation.
Umk 111 Die 2spp: 14 indivs; with cuticle.
Lit 111 Dic/Hei: 10 indivs; with cuticle, clear hairs.
Maz 21 1 Hei/Dic: 63 indivs; clear hairs & preservation.
Maz 111 Die era: 27 indivs; comparative full palaeodeme.
Specific diagnosis
A Fraxinopsis species of small size, with a wing showing ca 7 simple
to occasionally forking veins, and hirsute seed body with small auricles.
Specific characters
Seed: small (ca 12-21 mm long); wing partially hirsute, veins ca 7 at widest
point, occasionally forking, rarely anastomosing; seed body distinctly
hirsute, relatively large (V3— '/2 length of wing), with small lateral auricles.
Etymology
andium— the name given by Frenguelli (1941a), probably referring to the
Andes Mountains.
Comment & comparison
This species, as recorded from 16 Molteno TCs, is particularly vari-
able, both within and between palaeodemes. The seven best sampled of the
16 palaeodemes are illustrated here to show this intraspecific diversity.
Each palaeodeme is quite distinctive and each probably merits subspecific
status— some might merit species status— but the morphological distance
between them is far less than that which separates out the species F. auricu-
lata and F. cornicordis. The most diagnostic features of the seven palaeo-
demes are noted under the line illustrations.
PRE/F/ PRE/F/ PRE/F/
11740 4768a 11706
pi 145(1) pi 145(2) pi. 145(6)
Maz 211
PRE/F/ II 703
pi. 145(5)
veins 8 to 9, commonly forking to lateral and
distal margins, seed body and wing margins
distinctly hirsute.
pl. 145(4) PRE/F/4741
pi 145(3)
'rt all R2
3978 3981 3966V
Maz 111
W'O
PRE/F/
3985
PRE/F/
3984
wing relatively linear, veins commonly
anastomosing around distal margin
% allR2
Fraxinopsis
FRAXINOPSIALES
c/tRELITZIA 15 (2003)
375
Fraxinopsis auriculata J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/9408, pi. 147(5).
Assemblage: Wal 1 1 1 Die odo, Waldeck.
Preservation: virtually complete seed, no counterpart; impression in thickly
laminated, moderately baked, medium light grey shale with very good
cleavage.
Reference palaeodeme
Assemblage ( TC): as for holotype.
Specimens: 12 indivs; pi. 147(1-9).
Sister palaeodeme— nil.
Specific diagnosis
A Fraxinopsis species of medium size, with a wing showing ca 10
forking and anastomosing veins, and glabrous seed body with well-developed
lateral auricles.
Specific characters
Seed: medium (ca 32-A2 mm long); wing glabrous, veins ca 10 at widest
point, commonly forking, often anastomosing towards apex; seed body
glabrous, medium (’/4 length of wing), with well-developed lateral
auricles.
Etymology
auriculata— auricula (Lat.), ear; with reference to the diagnostic auricles.
Comment & comparison
F. auriculata , known only from the single palaeodeme (12 individuals)
from Waldeck (Wal 111), differs from the other species in the large pro-
truding auricles which are consistently developed. F. andium has much
smaller auricles.
The function of the auricles is not understood. They appear not to be
present in any other plant group with winged seeds.
Fraxinopsis cornicordis J.M.And. & H.M.And., sp. nov.
Holotype
Specimen: PRE/F/8225; pi. 148(2, 4, 6).
Assemblage: Kra 31 1 Die odo. Kraai River.
Preservation: virtually complete seed, part and counterpart; impression in
very thin-bedded, medium grey cherty shale with moderate cleavage.
Reference palaeodeme
Assemblage (TC): as for holotype.
Specimens: 35 indivs; pi. 148 (1-6).
Sister palaeodeme— nil.
Specific diagnosis
A Fraxinopsis species of large size, with a wing showing ca 16 frequently
anastomosing veins, and glabrous seed body without auricles.
Specific characters
Ovule! seed: large (up to >50 mm long); wing glabrous, veins ca 16 at
widest point, commonly forking, frequently anastomosing towards
apex; seed body glabrous, relatively small (‘/s length of wing), without
auricles.
Etymology
cornicordis— cornix (Lat.), crow; cordis (Lat.), heart; with reference to the
type locality and the shape of the seed body.
Comment & comparison
F. cornicordis is represented by the single palaeodeme of 35 individuals
from Kraai River (Kra 311). It is larger in size than F. auriculata, and differs
in the lack of auricles. It differs from the remaining species in its large size
and frequently anastomosing veins. Ganuza et al. (1995) illustrated two
specimens from Argentina which are tentatively placed in this species
based on their large size.
FRAXINOPSIALES
Fraxinopsis
376
c/tRELITZIA 15 (2003)
Yabeiella Oishi 1931
Type species
Yabeiella brackebuschiana (Kurtz ) Oishi 1931.
Cacheuta, N. Argentina; Camian, Triassic.
Generic concept
A stem-gnetopsid leaf of linear to narrowly elliptic shape, with entire
lamina and occasionally to frequently anastomosing taeniopteroid venation
ending at an intramarginal vein.
Generic characters
Leaf: small to medium, linear to narrowly elliptic; lamina undivided,
entire, tip roundly obtuse to retuse, base cuneate to attenuate, without
distinct petiole; venation taeniopteroid, closely to well spaced, extend-
ing at ca 20-30° from horizontal, forking to repeatedly anastomosing,
with intramarginal vein.
Cuticle', see And. & And. (1989, p. 472); and this vol., tfs 11, 12 opposite.
Eponymy
Yabiella— named by Oishi (1931) in honour of Prof. H. Yabe.
Global range: 2 spp.. Gondwana, Tr. (LAD-NOR).
First: Yabeiella sp. (Webb 1980); UQL.4255, Esk Fm., Ipswich/Esk,
Queensland, Australia.
Last: Yabeiella sp. (Webb 1980); UQL.4259, Woogaroo Fm., Brisbane,
Queensland, Australia.
Gondwana Triassic occurrence
Frequency (F): 2 1 degree squares (of the 84 across Gondwana).
Ubiquity (U): 3 continents (of 5 comprising Gondwana).
Diversity (D): 2 foliage species.
Abundance (A): <1% (the norm as in Molteno TCs).
Longevity (L): 17 myrs (early Ladinian to later Norian).
Colonisation success: FUDAL rating — 2 1/3/2/— /1 7 = 43.
Intermediate success (Grade 3): Yabeiella was the 10th most prominent
Gondwana Triassic genus; it was frequent, ubiquitous and relatively
long-lived, but of low diversity and abundance.
Endemism: both described species (Y. brackebuschiana and Y. mareyesiaca)
are widespread.
Molteno occurrence
Frequency (F): 29 TCs (of 100 sampled in the Molteno).
Diversity (D): 2 species.
Abundance (A): occasional (1-2%) in 5 TCs; rare to very rare (<1%) in the
other 24 TCs.
Habit: probably a medium-sized tree.
Preferred habitat: ubiquitous in Dicroidiwn riparian forest and closed
woodland of the lake margins; far less frequent in Dicroidium open
woodland ( 10 of 3 1 TCs).
Affiliated organs
Female strobilus: unknown.
Seed: Fraxinopsis— Grade 4 (Cut. cor., Mut. occ.).
Male strobilus: unknown.
Classification & comparison
Intergeneric comparison
Gondwana Triassic gymnosperm genera— Yabeiella has a number of
microfloral features in common with Jungites and Gontriglossa. They have
remarkably similar cuticle with amorphously shaped nonpapillate cells
with meandering walls. In megafloral features they are quite distinct:
Yabeiella having a marginal vein and few anastomoses, Gontriglossa
strongly anastomosing venation, and Jungites occasional anastomoses and
an incised lamina margin.
Other genera— We are unaware of any other genera with marginal
veins that might be confused with Yabeiella.
Interspecific comparison
If judged purely on the basis of their leaf macromorphology, the two
species of Yabeiella recognised here from the Gondwana Triassic might be
considered separate genera. However, the fair quality cuticle that has been
prepared from both species is more or less identical (And. & And. 1989).
In the Molteno and across Gondwana, the co-occurrence (affiliation)
between the leaf genus Yabeiella and the seed genus Fraxinopsis has been
firmly established; but on present evidence we are unable to propose links
at specific level.
The reference palaeodemes of the two species do not overlap. The full
set of 29 palaeodemes of Y. brackebuschiana and three of Y. mareyesiaca
from the Molteno Fm. show no intermediates between the two species,
although they occur together at Fut 2 1 1 , Umk 111 and Aas 411.
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
Yabeiella
FRAXINOPSIALES
dA-RELITZIA 15 (2003)
377
FRAXINOPSIALES
Yabeiella
378
<7j/tRELITZI A 15 (2003)
(Lit 111 Dic/Hei) g
mmmms
X100
BP/2/2200
I WWM
X250
BP/2/2200
BP/2/2200
PRE/F/22506
X250
Fraxinopsis andium
pi. 143
FRAXINOPSIALES
c/trELITZIA 15 (2003)
379
PRE/F/5324
PRE/F/ 10039
Birds River
(Bir 1 1 1 Sph 2spp)
BP/2/5305
BP/2/5316
PRE/F/10499 X4
PRE/F/11818 X4
BP/2/5305
PRE/F/19923
PRE/F/15613a
PRE/F/15613b
FRAXINOPSIALES
pi. 144
Fraxinopsis andium
380
d/TRELITZIA 15 (2003)
Fraxinopsis andium
pi. 145
FRAXINOPSIALES
c/tRELITZIA 15 (2003)
381
Sani Pass
(San 111 Die era)
PRE/F/18713
• ' \sC '1.
SS&
BP/2/2680
FRAXINOPSIALES
pi. 146
F raxinopsis andium
382
^TRELITZIA 15 (2003)
PRE/F/94 16
PRE/F/9418 'if. ' X2
PRE/F/94 19
PRE/F/9418
PRE/F/9414
Waldeck
(Wal 1 1 1 Die odo)
PRE/F/9408
saoBeexsuwaESfUaii
Holotype
v: 7^ n.
PRE/F/9418 •
PRE/F/94 16
PRE/F/9414
F raxinopsis auriculata
pi. 147
FRAXINOPSIALES
^/tRELITZIA 15 (2003)
383
FRAXINOPSIALES
pi. 148
Fraxinopsis cornicordis
384
g/trELITZIA 15 (2003)
Jungites J.M.And. & H.M.And. 1989
Type species
Jungites polymorpha J.M.And. & H.M.And. 1989.
Little Switzerland, Karoo Basin, S. Africa; Camian, Triassic.
Generic diagnosis
A stem-gnetopsid leaf of narrowly to broadly elliptic shape, with irregu-
larly cleft lamina and taeniopteroid venation with occasional anastomoses.
Generic characters
Attachment: fascicle of several leaves on (?)short shoot.
Leaf, small to relatively large, narrowly to (?)broadly elliptic; lamina irregu-
larly and variously cleft, margin very finely irregularly serrulate; tip
acute to obtuse; base cuneate to attenuate, without distinct petiole;
venation taeniopteroid, closely to well spaced, extending at low angle
from horizontal, occasionally forking and anastomosing.
Cuticle: see And. & And. (1989, p. 497); this vol., tfs 4, 6 opposite.
Eponymy
Jungites— for Carl Jung, Swiss psychiatrist with a particular interest in
schizophrenia (split personality). The diagnostic feature of the leaf
Jungites is the irregularly cleft lamina.
Global range: 3 spp., Gondwana, M.-U.Tr. (AWS-CRN).
First: J. reservoirensis And. & And. (1989), Burgersdorp Fm., Reservoir,
Aliwal North.
Last: J. polymorpha. Molteno Fm.
Gondwana Triassic occurrence
Frequency (F): 3 degree squares (of the 84 across Gondwana).
Ubiquity (U): 2 continents (of 5 comprising Gondwana).
Diversity (D): 3 foliage species.
Abundance (A): <1% (the norm in Molteno TCs).
Longevity (L): 13 myrs.
Colonisation success: FUDAL rating 3/2/3/-/13 = 21.
Limited success (Grade 2): Jungites was the 17th most prominent
genus in the Gondwana Triassic; it was relatively long-lived, but of
low frequency, ubiquity, diversity and abundance.
Endemism: all 3 species are single-assemblage endemics.
Molteno occurrence
Frequency (F): 1 TC (of 100 sampled in the Molteno).
Diversity (D): 2 species.
Abundance (A): very rare (<1%) in the single TC.
Affiliated organs: unknown.
Classification & comparison
Intergeneric comparisons
Gondwana Triassic gymnosperm genera— As noted earlier, Jungites has a
range of cuticular features suggesting an obvious relationship with
Yabeiella and Gontriglossa (for further comment on the similarities and
differences between the three genera, see text under Yabeiella , p. 376).
Other genera— Had Jungites not yielded cuticle, we might have included
the component species in the cycad genus Nilssonia or in one of the
cycadeoid genera Nilssoniopteris or Anomozamites. Nilssonia and Nilssoni-
opteris have entire to irregularly divided laminae similar to Jungites, while
some species of Nilssoniopteris and Anomozamites also resemble it with
their irregularly serrulate margins. The cuticle of Jungites, however, is quite
unlike that in any cycad or cycadeoid and separates it from those genera.
Interspecific comparisons
The three species of Jungites recognised here from the Gondwana
Triassic are perfectly distinct, though superficially similar in leaf macro-
morphology and fall reasonably within the compass of a natural genus. Fair
quality cuticle has been derived from both Molteno species, J. polymorpha
and J. sp.A, and proved to be virtually identical. The third species, J. reser-
voirensis, recorded from the early Middle Triassic Burgersdorp Fm. of the
Karoo Basin, is from a site without cuticle (And. & And. 1989). A further
species possibly occurs in South America (Menendez 1951), but the two
recorded individuals were included as sp. indet. in our earlier revision
(And. & And. 1989).
GONDWANA TRIASSIC, GEOSTRAT' DISTRIBUTION
Jungites
FRAXINOPSIALES
C^TRELITZIA 15 (2003)
385
Jungites
Lit ill
reference palaeodeme
Jungites spA
Lit 111
reference palaeodeme
FRAXINOPSIALES
Jungites
386
^/tRELITZIA 15 (2003)
Umkomaas Valley
(Umk 111 Die 2spp)
Little Switzerland
(Lit 1 1 1 Dic/Hei)
UNDESCRIBED SEEDS
(pis 149-152)
Waldeck
(Wal 1 1 1 Die odo)
Undescribed seeds
pi. 149
INCERTAE/INCERTAE
387
INCERTAE/INCERTAE
pi. 150
Undescribed seeds
d/TRELITZIA 15 (2003)
Aasvoelberg
(Aas 411 Dic/Sph)
r
§
388
(SP X
RELITZIA 15 (2003)
Undescribed seeds
pi. 151
INCERTAE/INCERTAE
389
^j/trELITZIA 15 (2003)
INCERTAE/INCERTAE
pi. 152
Undescribed seeds
Umkomaas Valley
(Umk 111 Die 2spp)
Aasvoelberg
(Aas 411 Dic/Sph)
PRE/F/22518
390
d^TRELITZIA 15 (2003)
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d^TRELITZIA 15 (2003)
GLOSSARY
The terms included here are those considered most relevant to our study of Molteno and Gondwana Triassic floristics
and biodiversity. They are arranged to emphasise the significance of certain fields. A few terms are not applied in this
book, but are retained for coherence of related topics. The glossaries appearing in the sequels to this work (Anderson
et al.. in prep.; And. & And., in prep.) will be more comprehensive in certain fields such as ‘Evolution and genetics’.
Morphological terminology, covered in a pictorial key on pp. 50-53, is not repeated here.
**— terms introduced in our own works on the Molteno or other fossil floras of South Africa.
*— usage followed in our own works.
SAMPLING
Optimal sampling of fossiliferous strata— and there are many criteria
that might be considered in defining optimal— is critical to reliable track-
ing of diversity patterns through time.
Frequency— The measure of frequency of a fossil taxon within a formation
is the number of taphocoenoses (TCs) or assemblages, out of the total
sampled, in which it has been found. Through the Gondwana Triassic,
for instance, it might be the number of degree squares (out of the total
sampled) in which it has been found. *
Abundance— The abundance of a fossil taxon is a measure of the absolute
or relative number of individuals collected from or encountered in an
assemblage, formation, region or continent. *
Assemblage— The full suite of fossil individuals or palaeodemes collected
from a distinct lithological unit (lithosome) of limited geographic and
stratigraphic extent. A megaplant assemblage will generally represent
a localised mosaic of plant associations, less often a single association,
through several generations. *
Locality— An area to about 1 km in diameter, which may include continu-
ous fossiliferous exposure, but will generally include one or more pro-
ductive exposures of lesser rank. *
Extensive sampling— Concerns the number and spread of localities sampled
in a fossiliferous horizon. *
Intensive sampling— Concerns the comprehensiveness of collecting from a
particular fossiliferous deposit. *
Singletons— Those species recorded from a particular geological horizon
(member, formation or biozone) that are known from only a single
locality. The number of singletons (and doubletons) is critical in pro-
jecting biodiversity from the observed record.
Doubletons— Those species recorded from a particular geological horizon
that are known from only two localities.
TAPHONOMY
There is inevitably a reduction between the number of species making
up the original flora and fauna of the region and that preserved in the fossil
record. This filtering or winnowing of the original diversity occurs through
the taphonomic process. Taphonomy is of vital concern in biodiversity pro-
jections. (Selected references: Ferguson 1992; Behrensmeyer et <j/.1992,
Cuneo et al. 1993.)
Taphonomy— The conditions and processes intervening between a living
species association and a fossil assemblage.
Taphonomic mode— Reflects the environmental context of the fossiliferous
deposit. Examples include levee, lake, crevasse splay or abandoned
channel of the floodplain (i.e. subenvironments of meandering and
braided fluvial systems).
Isotaphonomic— Refers to fossiliferous assemblages of like taphonomic
mode.
Biocoenosis— Living community, (p. 25)
Thanatocoenosis— Death assemblage; that fraction of the original commu-
nity that is preserved in the fossil deposit, (p. 25)
Taphocoenosis (TC)— Taphonomic assemblage; fossil assemblage result-
ing from the taphonomic process; the aggregate of fossil remains con-
tained in a deposit or bed.
Phytotaphocoenosis— A fossil plant assemblage or plant taphocoenosis.
Autochthonous— Describes TCs that have originated in situ. While such
assemblages might be relatively common in the case of fossil
tetrapods, they are a far more rare phenomenon in palaeobotany (e.g.
horsetails entombed in growth position in a marsh).
Parautochthonous— Describes TCs deposited close to the original site of
occurrence.
Allochthonous— Describes TCs deposited at some significant distance from
the original site of occurrence.
PALAEODEMES
Terms either from And. & And. (1983) or our later works.
Palaeodeme (fossil population)— A collection of fossil specimens judged
to represent a single potentially interbreeding population of plants or
animals, showing a normal distribution of variation for selected diag-
nostic characters, and derived from a single taphocoenosis from a dis-
crete small-scale lithological unit (lithosome) such as an abandoned
channel infill or crevasse splay. **
Reference palaeodeme (RP)— The most comprehensive, representative,
photographically documented palaeodeme in the literature proposed as
reference for a particular infrageneric taxon. **
Reference assemblage (RA)— That assemblage from which the reference
palaeodeme derives. **
Reference specimen (RS) — The most complete, average, mature specimen
selected from the reference palaeodeme. **
Home palaeodeme— The palaeodeme from which a specimen derives. **
Sister palaeodeme— A palaeodeme belonging to the same species. **
FOSSIL TAXA
For form-genera and organ-genera, we follow standard usage. Further
terms in this field are introduced, or used in a particular sense, to meet the
specific needs of our Molteno research. The concepts can equally be
applied to species.
Form-genus—' a genus of fossil plants based on a detached organ which,
because of the limited characters shown, cannot be assigned to a fam-
ily, although it may be possible to assign it to a higher taxonomic level.
The “artificial” nature of such a genus is illustrated by the several
instances where we have one form-genus in which different species
may closely resemble members of two or more different families’
(Jones & Rowe 1999).
Organ-genus—' a genus based on only part (an organ or organs) of a fossil
plant, showing a sufficient range of distinctive characters that it may
reasonably be assigned to a family. An organ-genus is regarded as “nat-
ural”, in the sense that its constituent species are believed to have the
same close relationship as those of a living genus. However, an organ-
genus differs from a genus of living plants in that only fossils of the
same organ, showing the same type of characters, can be assigned to it
(e.g. fossil lauraceous leaves cannot be assigned to Laurocarpum, an
organ-genus of fossil lauraceous fruit)’ (Jones & Rowe 1999).
Whole-plant genus— A fossil-plant genus considered ‘natural’ that includes
one or more organ-genera. The term is applied here only after com-
prehensive and systematic affiliation studies for the group (e.g. gym-
nosperms) and formation (e.g. Molteno) have been completed. In the
Molteno (see Tab. 15, p. 21), we recognise 38 gymnosperm whole-
plant genera, i.e. 38 genera known from at least one of the three
organs— ovulate, foliage or microsporangiate. (p. 43) *
Fossil-plant genus (or simply plant-genus)— A generalised term usually
referring to a ‘natural’ whole-plant or multi-organ genus. *
Multi-organ genus— A fossil-plant genus considered ‘natural’ that includes
more than one organ-genus. In the Molteno (see Tab. 15, p. 21), we
recognise 16 gymnosperm multi-organ genera, i.e. 16 genera known
from at least two of the three organs— ovulate, foliage or microspo-
rangiate. (p. 43) *
Mother-plant genus— The original living-plant genus occurring in the
Molteno Biome. (The relationship between the living and fossil genus
is akin to that between the phytocoenosis and taphocoenosis, i.e.
between the living assemblage and the fossil assemblage.) *
Glossary
(^/trelitzia 15 (2003)
397
Sister genera— Those organ-genera or whole-plant genera included in the
same family or order (e.g. the three male-cone genera Fredianthus ,
Lutanthus and Odyssianthus , from the Molteno placed in the order
Voltziales, pp. 70, 74). *
Sister specimen— A specimen of the same species from the same
palaeodeme (e.g. as for Lutanthus hemidiscus and L. ornatus, p. 76). *
BIODIVERSITY
‘The variety of organisms considered at all levels, from genetic vari-
ants belonging to the same species through arrays of species to arrays of
genera, families, and still higher taxonomic levels; includes the variety of
ecosystems, which comprise both the communities of organisms within
particular habitats and the physical conditions under which they live’
(Wilson 1992).
Observed diversity— The actual tally of taxa of a particular rank (e.g.
species, families, orders) collected (curated and/or described) from a
particular geological horizon (e.g. formation, assemblage zone). *
Preserved diversity— The projected tally of taxa of a particular rank repre-
senting the full potential sample (assuming comprehensive sampling
of all preserved taphocoenoses) from a particular geological horizon. *
Existed diversity— The projected tally of taxa of a particular rank repre-
senting the full flora or fauna that actually inhabited the various habi-
tats of the biome existing at the time of deposition of a particular geo-
logical horizon. *
Alpha diversity— The number of species occupying a particular habitat at a
particular locality, e.g. the 73 foliage species recorded from the ripari-
an forest at the Umkomaas locality, Molteno Formation.
Beta diversity— The rate at which the species number increases along a
transect through adjacent ecozones (habitats).
Gamma diversity— The totality of species, considering all habitats over a
broad area; e.g. the full extent of the Molteno Biome.
MNS (Minimum number of species)— The minimum tally of observed
whole-plant species in a taphocoenosis, formation or other defined
body of sediment. In the Molteno (Tab 15, p. 21), we recognise 143
gymnosperm whole-plant species; this is the sum of MNSs recognised
for each whole-plant genus in the formation. *
ECOLOGY & PALEOECOLOGY
Biome— ‘A major category of habitat in a particular region of the world,
such as the tundra of northern Canada or the rainforest of the Amazon
basin’ (Wilson 1992).
Ecosystem — ‘The organisms living in a particular environment, such as a
lake or a forest (or, in increasing scale, an ocean or the whole planet),
and the physical part of the environment that impinges on them. The
organisms alone are called the community’ (Wilson 1992).
Ecozone (habitat)— A physical or vegetational environment of a particular
restricted kind, such as the riparian forests, braided-river sandbar
meadows or floodplain woodlands of the Molteno Biome.
Guild— A set of species that live within a community in the same area and
harvest the same food by similar means (Wilson 1992).
EVOLUTION & GENETICS
‘Evolution proceeds mostly by the accidental substitution of one or
more of the letters (of the genetic code), followed by the winnowing of
these mutations and their combinations through natural selection' (Wilson
1992). The immense diversity of species derives from the astronomic num-
ber of rearrangements possible within the hugely lengthy sequence of
nucleotide letters of the genetic code.
Evolutionary biology— Covers the broad array of disciplines— palaeontol-
ogy, ecology, population biology, systematics, biogeography, ethology,
cladistics, molecular evolution etc.— focused on the evolutionary
process and consequently the building of biodiversity.
Hox genes— Those genes that control (encode) the body plans of embryos
of all animals, apparently from the first multicellular animals some 700
Ma back to fruit flies, elephants and human beings today. All mam-
mals. for instance, have 38 different Hox genes.
Genetic code— The code contains ca 1 million nucleotide pairs in bacteria
and 1 to 10 billion pairs in higher plants and animals (Wilson 1992).
Macroevolution— ‘Evolution above the species level’ (Stanley 1979).
Microevolution— Evolutionary change within the species (Stanley 1979).
Allopatric speciation (geographic speciation)— ‘The divergence to species
level by populations that originally belonged to the same species but
were isolated by a physical barrier such as a sea strait, river valley, or
mountain range’ (Wilson 1992).
Parapatric speciation— The divergence to species level by a local popula-
tion while remaining in general contact with other populations of the
original species (Stanley 1979).
Phyletic gradualism— Sympatric speciation. The evolution of one species
into another through time. One species replaces another through a
series of subspecies in an evolutionary lineage. Modification occurs
through geological time in a single phyletic line (phylogenesis).
(This and the next two terms from And. & And. 1983, after Sylvester-
Bradley 1977.)
Punctuated equilibria — Allopatric (geographic) speciation. A single
species becomes ancestral to two or more descendant species through
the splitting of the phyletic line (cladogenesis). Speciation occurs rel-
atively rapidly such that transitional forms will rarely be preserved in
the fossil record.
Reticulate speciation— Combines aspects of the previous two, involving
both phylogenesis and cladogenesis. The central characteristic is the
complex alternating isolation and hybridisation of races.
Extinction— The termination of any lineage of organisms, from subspecies
and species to classes and phyla (Wilson 1992).
Renewal— Refers here to the re-establishment of diversity after a mass global
extinction.
Adaptive radiation— 'The rapid proliferation of new taxa from a single
ancestral group’ (Stanley 1979).
Evolutionary success (or prominence)— Can be measured in many ways,
but is taken here as a combination of frequency, ubiquity, diversity,
abundance and longevity —giving a FUDAL rating (And. & And. 1999).
Frequency— measure of repetitiveness of occurrence.
Ubiquity— measure of general range of occurrence.
Diversity— measure of speciation, radiation, variability.
Abundance— measure of quantity.
Longevity— measure of duration of the lineage.
In the Molteno Formation, for instance, Dicroidium, with a FUDAL
rating of 188, wa9 clearly the most successful plant genus, as it was
throughout the Gondwana Triassic (pp. 26-29). *
CLADISTICS
Aside from its rigour in revealing phylogenetic relationships, cladistics
has a major role to play in biodiversity studies. Consider current knowledge
of mammalian and lepidopteran phylogeny: the cladogram. in each case,
brings into focus a considerable hidden, diversity during the earlier phases
of adaptive radiation of the group (And. 1999).
Cladistics— Phylogenetic taxonomy; a rigorous attempt— with each step
subject to Popperian falsification— to construct phylogenetic trees
based on a selection of morphological and/or molecular characters
with polarised (primitive or derived) states.
Phylogeny— The evolutionary history of a specified group of organisms of
any rank, with particular focus on the genealogical tree of lineages
comprising the group.
Cladogram— Figure or phylogenetic tree showing the branching pattern of
the group under study.
Lineage— ‘ A single line of descent’ (Stanley 1979).
Clade—A cluster of lineages deriving from a common ancestor.
Monophyletic — Pertaining to a group of taxa sharing a common ancestor.
Polyphyletic— Pertaining to a group of diverse taxa not sharing a common
ancestor.
Crown group— Includes the cluster of lineages expected j show all apo-
morphies (derived/advanced characters) of the extant taxa.
Stem group— Includes those forms (directly on the line to the extant group
or on extinct side-branches) showing only some of the apomorphies of
the extant taxa.
Sister group— That taxon or clade judged to share an immediate common
ancestor with the group in question.
Plesiomorphic— Pertaining to the primitive state of a character.
Apomorphic— Pertaining to the derived state of a character.
Glossary
398
d/TRELITZIA 15 (2003)
INDEX TO CURRENT MOLTENO GENERA
The index includes only the generic names— ovulate, microsporangiate and foliage— retained in this volume for the
gymnosperms of the Molteno Formation.
Alexia 318
Androstrobus 136
Antevsia 154
Avatia 192
Avistrobus 130
Bariopteris 324
Cetifructus 314
Cetiglossa 369
Clariphyllum 100
Ctenis 144
Cycadolepis 341
Dejerseya 186
Dicroidium 256
Dordrechtites 60
Eosteria 194
Fanerotheca 272
Fraxinopsis 370
Fredianthus 70
Fredlindia 336
Ginkgoites 198
Gontriglossa 364
Graciliglossa 368
Gypsistrobus 126
Flalleyoctenis 344
Flamshawvia 210
Heidiphyllum 90
Flelvetianthus 132
Hlatimbia 322
Hystricia 330
Jeanjacquesia 142
Jungites 384
Kannaskoppia 286
Kannaskoppianthus 290
Kannaskoppifolia 294
Kurtziana 176
Leguminanthus 342
Lepidopteris 156
Lindtheca 356
Linguifolium 334
Lutanthus 74
Matatiella 172
Moltenia 146
Nataligma 362
Odyssianthus 88
Pagiophyllum 124
Paraginkgo 208
Peltaspermum 148
Pseudoctenis 140
Pteruchus 250
Rissikia 112
Rissikianthus 108
Rissikistrobus 102
Saportaea 332
Scytophyllum 170
Sphenobaiera 222
Stachyopitys 216
Switzianthus 182
Taeniopteris 358
Telemachus 82
Umkomasia 240
Weltrichia 340
Yabeiella 376
Index
STRELITZIA
1. Botanical diversity in southern Africa. 1994. B.J. Huntley (ed.). ISBN 1-874907-25-0.
2. Cyperaceae in Natal. 1995. K.D. Gordon-Gray. ISBN 1-874907-04-8.
3. Cederberg vegetation and flora. 1996. H.C. Taylor. ISBN 1-874907-28-5.
4. Red Data List of southern African plants. 1996. Craig Hilton-Taylor. ISBN 1-874907-29-3.
5. Taxonomic literature of southern African plants. 1997. N.L. Meyer, M. Mossmer & G.F. Smith (eds). ISBN
1-874907-35-8.
6. Plants of the northern provinces of South Africa: keys and diagnostic characters. 1997. E. Retief & P.P.J. Herman. ISBN
1-874907-30-7.
7. Preparing herbarium specimens. 1999. Lyn Fish. ISBN 1-919795-38-3.
8. Bulbinella in South Africa. 1999. Pauline L. Perry. ISBN 1-919795-46-4. OUT OF PRINT.
9. Cape plants. A conspectus of the Cape flora of South Africa. 2000. P. Goldblatt & J.C. Manning. ISBN 0-620-26236-2.
10. Seed plants of southern Africa: families and genera. 2000. O.A. Leistner (ed.). ISBN 1-919795-51-0.
11. The Cape genus Lachnaea (Thymelaeaceae): a monograph. 2001. J.B.P. Beyers. ISBN 1-919795-52-9.
12. The Global Taxonomy Initiative: documenting the biodiversity of Africa/LTnitiative Taxonomique Mondiale: documenter
la biodiversite en Afrique. R.R. Klopper, G.F. Smith & A.C. Chikuni (eds). 2001. ISBN 1-91 9795-63-4. OUT OF PRINT.
13. Medicinal and magical plants of southern Africa: an annotated checklist. 2002. T.H. Arnold. C.A. Prentice, L.C. Hawker,
E.E. Snyman, M. Tomalin, N.R. Crouch & C. Pottas-Bircher. ISBN 1-919795-62-6.
14. Plants of southern Africa: an annotated checklist. 2003. G. Germishuizen & N.L. Meyer (eds). ISBN 1-919795-99-5.
15. Heyday of the gymnosperms: systematics and biodiversity of the Late Triassic Molteno fructifications. 2003. J.M. Anderson
& H.M. Anderson. ISBN 1-919795-98-7.
MEMOIRS OF THE BOTANICAL SURVEY OF SOUTH AFRICA
(discontinued after No. 63)
Still available:
2. Botanical survey of Natal and Zululand. 1921. R.D. Aitken & G.W. Gale.
8. Researches on the vegetation of Natal. Series II. 1925. J.W. Bews & R.D. Aitken.
17. The vegetation of the Divisions of Albany and Bathurst. 1937. R.A. Dyer.
29. The wheel-point method of survey and measurement of semi-open grasslands and karoo vegetation in South Africa. 1955.
C.E.M. Tidmarsh & C.M. Havenga.
31. Studies of the vegetation of parts of the Bloemfontein and Brandfort Districts. 1958. J.W.C. Mostert.
33. The vegetation of the Districts of East London and King William’s Town, Cape Province. 1962. D.M. Comins.
39. Flora of Natal. 1973. J.H. Ross. ISBN 0-621-00327-1.
41. The biostratigraphy of the Permian and Triassic. Part 3. A review of Gondwana Permian palynology with particular refer-
ence to the northern Karoo Basin, South Africa. 1977. J.M. Anderson. ISBN 0-621-03834-2.
42. Vegetation of Westfalia Estate on the north-eastern Transvaal escarpment. 1977. J.C. Scheepers. ISBN 0-62 1 -03844-X.
43. The bryophytes of southern Africa. An annotated checklist. 1979. R.E. Magill & E.A. Schelpe. ISBN 0-621 -047 18-X.
44. A conspectus of the African Acacia species. 1979. J.H. Ross. ISBN 0-621-05309-0.
45. The plant ecology of the Isipingo Beach area. Natal, South Africa. 1980. C.J. Ward. ISBN 0-621-05307-4.
46. A phytosociological study of the Upper Orange River Valley. 1980. M.J.A. Werger. ISBN 0-621-05308-2.
47. A catalogue of South African green, brown and red algae. 1984. S.C. Seagrief. ISBN 0-621-07971-5.
49. Pattern analysis in savanna-woodlands at Nylsvley, South Africa. 1984. R.H. Whittaker, J.W. Morris & D. Goodman. ISBN
0- 621-08265-1.
50. A classification of the mountain vegetation of the Fynbos Biome. 1985. B.M. Campbell. ISBN 0-621-08862-5.
52. A plant ecological bibliography and thesaurus for southern Africa up to 1975. 1986. A.P. Backer, D.J.B. Killick & D.
Edwards. ISBN 0-621-08871-4.
53. A catalogue of problem plants in southern Africa, incorporating the National Weed List of South Africa. 1986. M.J. Wells,
A.A. Balsinhas, H. Joffe, V.M. Engelbrecht, G. Harding & C.H. Stirton. ISBN 0-621-09688-1.
55. Barrier plants of southern Africa. 1987. L. Henderson. ISBN 0-621-10338-1.
57. Veld types of South Africa 3rd edn. 1988. J.P.H. Acocks. With separate wall map. ISBN 0-621-1 1394-8.
59. Tannin-like substances in grass leaves. 1990. R.P. Ellis. ISBN 0-620-15 15 1-X.
60. Atlas of the leaf anatomy in Pentaschistis (Arundineae: Poaceae). 1992. R.P. Ellis & H.P Linder. ISBN 0-9583205-1-9.
61. The marine red algae of Natal, South Africa: Order Gelidiales (Rhodophyta). 1992. Richard E. Norris. ISBN
1- 874907-01-3.
63. Biomes of southern Africa: an objective categorization. 2nd edn. 1994. M.C. Rutherford & R.H. Westfall. ISBN
1-874907-24-2.
ANNALS OF KIRSTENBOSCH BOTANIC GARDENS
(discontinued after Vol. 19)
The following volumes are available:
14. The moraeas of southern Africa. 1986. P. Goldblatt. ISSN 0-258-3305. ISBN 0-620-09974-7.
15. The botany of the southern Natal Drakensberg. 1987. O.M. Hilliard & B.L. Burtt. ISSN 0-258-3305. ISBN
0-620-10625-5.
18. The way to Kirstenbosch. 1988. D.P. McCracken & E.M. McCracken. ISSN 0-258-3305. ISBN 0-620-1 1648-X.
ENQUIRIES:
Bookshop, National Botanical Institute, Private Bag X101, Pretoria, 0001 South Africa.
Tel. (012) 804-3200 Fax (012) 804-321 1 E-mail bookshop@nbipre.nbi.ac.za http://www.nbi.ac.za
"This remarkable book on the gymnosperms of the Late
Triassic Molteno flora is a major contribution to
understanding the diversity of seed plants that existed in the
past. It joins the earlier works by the same authors in
reflecting two lifetimes dedicated to exploring the
biodiversity of the past so as to better illuminate the
biodiversity of the present. ... I am not aware of any other set
of fossil assemblages that has been collected with such
intensity, such uniformity of approach and such care. ... By
any measure, John and Heidi Anderson's dedication to
unlocking the secrets of the Molteno Formation has been
extraordinary and the result has been a landmark
contribution to palaeontology." Peter R. Crane, FRS,
Director, Royal Botanic Gardens, Kew, London, UK.
"The book is impressive in its scope and detail. It will
represent an invaluable contribution to the palaeobotanical
literature and will complete the description of the
gymnospermous components of the Upper Triassic Molteno
Formation — now the best studied Triassic flora on Earth. No
previous palaeobotanical studies have provided such a
comprehensive evaluation of the fossil-organ relationships
between ovuliferous fruits, microsporangiate fruits and
foliage from a formation covering such a large area and
represented by around 1 00 fossil assemblages and around
300 000 specimens." Stephen McLoughlin, University of
Melbourne, Australia.
"The prior volumes on fossil leaves have been invaluable,
and set a new standard for thoroughness of documentation.
. . . Few other publications can match their work as a
scientific resource. . . . This next volume is arguably the most
important of all as illustration of the reproductive organs will
reveal the biological affinities of the plants. . . . It will form a
fitting companion to the volumes already published and
represents another monumental contribution." Gregory J.
Retallack, University of Oregon, USA.
"It was 35 years ago, in March 1 967, that I ventured for the
first time into Molteno territory. From then till now the world
has changed. It has shifted — dramatically — in virtually every
respect: . . . Most importantly of all, from our perspective, the
human population has doubled from 3 billion to 6 billion
persons in this time . . . The Molteno became, increasingly, a
lot more than merely an exercise in collecting and describing
fossils. ... It became part of a multifronted obsession to help
swell our awareness of biodiversity trends through time, and
to impact somehow on mankind's collision with that diversity
as it exists today." John M. Anderson (from Preface).
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